<?xml version="1.0" encoding="UTF-8" standalone="no"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD Journal Publishing DTD v2.3 20070202//EN" "journalpublishing.dtd">
<article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" article-type="research-article">
<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Psychol.</journal-id>
<journal-title>Frontiers in Psychology</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Psychol.</abbrev-journal-title>
<issn pub-type="epub">1664-1078</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fpsyg.2018.02300</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Psychology</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>The Structural Effects of Modality on the Rise of Symbolic Language: A Rebuttal of Evolutionary Accounts and a Laboratory Demonstration</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name><surname>Boucher</surname> <given-names>Victor J.</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>&#x002A;</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/159602/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Gilbert</surname> <given-names>Annie C.</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/103360/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Rossier-Bisaillon</surname> <given-names>Antonin</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/524358/overview"/>
</contrib>
</contrib-group>
<aff id="aff1"><sup>1</sup><institution>Laboratoire de Sciences Phon&#x00E9;tiques, D&#x00E9;partement de Linguistique, Universit&#x00E9; de Montr&#x00E9;al</institution>, <addr-line>Montreal, QC</addr-line>, <country>Canada</country></aff>
<aff id="aff2"><sup>2</sup><institution>School of Communication Sciences and Disorders, McGill University</institution>, <addr-line>Montreal, QC</addr-line>, <country>Canada</country></aff>
<aff id="aff3"><sup>3</sup><institution>Centre for Research on Brain, Language and Music</institution>, <addr-line>Montreal, QC</addr-line>, <country>Canada</country></aff>
<author-notes>
<fn fn-type="edited-by"><p>Edited by: Jay Rueckl, University of Connecticut, United States</p></fn>
<fn fn-type="edited-by"><p>Reviewed by: Lorenzo Magrassi, University of Pavia, Italy; Alexander Bergs, Osnabr&#x00FC;ck University, Germany</p></fn>
<corresp id="c001">&#x002A;Correspondence: Victor J. Boucher, <email>victor.boucher@umontreal.ca</email></corresp>
<fn fn-type="other" id="fn002"><p>This article was submitted to Language Sciences, a section of the journal Frontiers in Psychology</p></fn>
</author-notes>
<pub-date pub-type="epub">
<day>28</day>
<month>11</month>
<year>2018</year>
</pub-date>
<pub-date pub-type="collection">
<year>2018</year>
</pub-date>
<volume>9</volume>
<elocation-id>2300</elocation-id>
<history>
<date date-type="received">
<day>25</day>
<month>08</month>
<year>2017</year>
</date>
<date date-type="accepted">
<day>05</day>
<month>11</month>
<year>2018</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#x00A9; 2018 Boucher, Gilbert and Rossier-Bisaillon.</copyright-statement>
<copyright-year>2018</copyright-year>
<copyright-holder>Boucher, Gilbert and Rossier-Bisaillon</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/"><p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p></license>
</permissions>
<abstract>
<p>Why does symbolic communication in humans develop primarily in an oral medium, and how do theories of language origin explain this? Non-human primates, despite their ability to learn and use symbolic signs, do not develop symbols as in oral language. This partly owes to the lack of a direct cortico-motoneuron control of vocalizations in these species compared to humans. Yet such modality-related factors that can impinge on the rise of symbolic language are interpreted differently in two types of evolutionary storylines. (1) Some theories posit that symbolic language originated in a gestural modality, as in &#x201C;sign languages.&#x201D; However, this overlooks work on emerging sign and spoken languages showing that gestures and speech shape signs differently. (2) In modality-dependent theories, some emphasize the role of iconic sounds, though these lack the efficiency of arbitrary symbols. Other theorists suggest that ontogenesis serves to identify human-specific mechanisms underlying an evolutionary shift from pitch varying to orally modulated vocalizations (babble). This shift creates numerous oral features that can support efficient symbolic associations. We illustrate this principle using a sound-picture association task with 40 learners who hear words in an unfamiliar language (Mandarin) with and without a filtering of oral features. Symbolic associations arise more rapidly and accurately for sounds containing oral features compared to sounds bearing only pitch features, an effect also reported in experiments with infants. The results imply that, beyond a competence to learn and use symbols, the rise of symbolic language rests on the types of signs that a modality of expression affords.</p>
</abstract>
<kwd-group>
<kwd>language evolution</kwd>
<kwd>symbolic communication</kwd>
<kwd>neurophysiology of speech</kwd>
<kwd>language development</kwd>
<kwd>comparative physiology</kwd>
</kwd-group>
<contract-num rid="cn001">no. 175811</contract-num>
<contract-sponsor id="cn001">Fonds Qu&#x00E9;b&#x00E9;cois de la Recherche sur la Nature et les Technologies<named-content content-type="fundref-id">10.13039/501100003150</named-content></contract-sponsor>
<counts>
<fig-count count="3"/>
<table-count count="0"/>
<equation-count count="0"/>
<ref-count count="173"/>
<page-count count="13"/>
<word-count count="0"/>
</counts>
</article-meta>
</front>
<body>
<sec><title>Introduction</title>
<p>There is a vast literature on the origin of spoken language, much of which offers diverging viewpoints with few areas of consensus. For instance, there is no agreement in this literature on how to define &#x201C;language&#x201D; (<xref ref-type="bibr" rid="B28">Christiansen and Kirby, 2003</xref>). On the other hand, it is widely accepted that a fundamental feature of language is its symbolic function and that, aside from humans, no other species have developed systems of signs such as those that appear in spoken language. Indeed, for some, humans are the <italic>symbolic species</italic> (<xref ref-type="bibr" rid="B39">Deacon, 1997</xref>). However, claiming the human specificity of symbolic communication rests on how one defines symbols, and the processes by which they evolved. The present paper aims at clarifying these processes within evolutionary theories while offering a demonstration of how the ability to articulate sounds presents an essential factor in the rise of symbolic language.</p>
<p>For some readers, this ability relating to a specific modality of expression may seem to be an obvious factor in the rise of symbolic communication. Yet general definitions of symbols often overlook processes of expression and how they contribute to the formation of signs. In fact, many evolutionists refer to <xref ref-type="bibr" rid="B144">Saussure (1916/1966)</xref> and <xref ref-type="bibr" rid="B123">Peirce (1998)</xref> and define &#x201C;symbols&#x201D; principally as arbitrary associations between signals and concepts of objects or events (i.e., &#x201C;referents&#x201D;). Authors also recognize that symbolic associations can operate from memory, when designated referents are not in the context of communication, a feature that <xref ref-type="bibr" rid="B65">Hockett (1960)</xref> called &#x201C;displacement.&#x201D; These criteria are useful in distinguishing symbols from signs that operate as <italic>icons</italic> or <italic>indices</italic>. The latter involve a <italic>non-arbitrary</italic> resemblance or physical connections to referents, whereas nothing in the attributes of symbols provides a clue as to their interpretation (<xref ref-type="bibr" rid="B41">Deacon, 2012</xref>, p. 14; <xref ref-type="bibr" rid="B159">Stjernfelt, 2012</xref>). However, definitions from Saussure and Pierce which focus only on arbitrary association can lead to a conceptualization of symbols as mental constructs, unrelated to modalities of expression. Indeed, Saussure saw language as reflecting a separate mental capacity or &#x201C;faculty&#x201D; that could generate symbols in <italic>any</italic> modality such as speech or gestures (<xref ref-type="bibr" rid="B144">Saussure, 1916/1966</xref>, pp. 10&#x2013;11). Such ideas have had a lasting influence, especially on linguistic theory, where language is seen to reflect a mental competence that has little to do with modalities of performance (<xref ref-type="bibr" rid="B21">Chomsky, 1965</xref>; <xref ref-type="bibr" rid="B59">Hauser et al., 2002</xref>). But if this is the case, then why is it that symbolic language develops primarily in a vocal medium?</p>
<p>In focusing on this question, the following discussion draws attention to a body of work in primatology which has failed to uncover a distinct mental ability that could account for symbolic language in humans. On the other hand, we outline that humans are the only primates that possess a cortical control of vocal signals so that, overall, the data undermine the belief that symbolic language arose from an amodal mental competence. A review of this belief that underlies popular theories of the origin of spoken language serves as background to a demonstration of an opposing modality-dependent principle where symbolic language is seen as relating to an ability to articulate sounds. Such a demonstration reflects the approach of a group of studies where evolutionary scenarios are submitted to critical laboratory experiments and computer simulations (as in <xref ref-type="bibr" rid="B56">Gasser, 2004</xref>; <xref ref-type="bibr" rid="B109">Monaghan and Christiansen, 2006</xref>; <xref ref-type="bibr" rid="B121">Oudeyer and Kaplan, 2006</xref>; <xref ref-type="bibr" rid="B172">Williams et al., 2008</xref>; <xref ref-type="bibr" rid="B110">Monaghan et al., 2011</xref>, <xref ref-type="bibr" rid="B111">2012</xref>, <xref ref-type="bibr" rid="B112">2014</xref>).</p>
<sec><title>Cognitive Skills as Insufficient Factors in the Rise of Symbolic Communication</title>
<p>In reviewing hypotheses of the origin of spoken language, it is important to acknowledge that several cognitive abilities and neural processes which were thought to underlie symbolic communication in humans have since been observed in other primates. In particular, it has been established that, with training, apes can learn vast sets of visual symbols and can combine these productively (e.g., <xref ref-type="bibr" rid="B147">Savage-Rumbaugh et al., 1993</xref>; <xref ref-type="bibr" rid="B150">Savage-Rumbaugh and Lewin, 1996</xref>; <xref ref-type="bibr" rid="B94">Lyn and Savage-Rumbaugh, 2000</xref>; <xref ref-type="bibr" rid="B145">Savage-Rumbaugh and Fields, 2000</xref>). Follow-up studies have documented that chimpanzees and bonobos raised in symbol-rich environments can develop a vocabulary and utterance complexity similar to those of 3 year-old children (<xref ref-type="bibr" rid="B82">Lieberman, 1984</xref>, chapter 10; <xref ref-type="bibr" rid="B54">Gardner and Gardner, 1985</xref>, <xref ref-type="bibr" rid="B55">1969</xref>; <xref ref-type="bibr" rid="B122">Pedersen, 2012</xref>). There are also reported cases where chimpanzees acquired elements of <italic>American Sign Language</italic> (ASL) only by communicating with other ASL-trained chimpanzees (<xref ref-type="bibr" rid="B54">Gardner and Gardner, 1985</xref>). Moreover, brain-imaging research indicates that associative memory in symbol learning involves similar neurological structures in apes, monkeys, and humans (e.g., <xref ref-type="bibr" rid="B158">Squire and Zola, 1996</xref>; <xref ref-type="bibr" rid="B44">Eichenbaum and Cohen, 2001</xref>; <xref ref-type="bibr" rid="B173">Wirth et al., 2003</xref>).</p>
<p>Other symbol-related abilities extend to non-human primates despite continuing claims to the contrary. Of note, the capacity to create hierarchical or embedded combinations of signs &#x2013; said to reflect a process of &#x201C;recursion&#x201D; &#x2013; was held to be uniquely human (<xref ref-type="bibr" rid="B59">Hauser et al., 2002</xref>; <xref ref-type="bibr" rid="B51">Fitch et al., 2005</xref>; cf. <xref ref-type="bibr" rid="B58">Hauser, 2016</xref>). Some also maintained that a related ability to combine symbols based on conceptual relations, a property termed &#x201C;Merge,&#x201D; was distinctly human (<xref ref-type="bibr" rid="B22">Chomsky, 1972</xref>, <xref ref-type="bibr" rid="B26">2012</xref>; <xref ref-type="bibr" rid="B14">Bolhuis et al., 2012</xref>; cf. <xref ref-type="bibr" rid="B88">Lieberman, 2016</xref>). However, <xref ref-type="bibr" rid="B127">Perruchet and Rey (2005)</xref> demonstrated that chimpanzees can learn to generate embedded sequences of given symbols (and see <xref ref-type="bibr" rid="B128">Perruchet et al., 2004</xref>, <xref ref-type="bibr" rid="B126">2006</xref>; <xref ref-type="bibr" rid="B157">Sonnweber et al., 2015</xref>). Additionally, research has shown that monkeys can distinguish acoustic cues in speech (as discussed by <xref ref-type="bibr" rid="B11">Belin, 2006</xref>), and manifest a &#x201C;statistical learning&#x201D; of speech sounds (<xref ref-type="bibr" rid="B60">Hauser et al., 2001</xref>). Several reports have further shown that non-human primates can process combinations of symbols based on conceptual relations (contra, e.g., <xref ref-type="bibr" rid="B68">Jackendoff, 2002</xref>; <xref ref-type="bibr" rid="B131">Pinker and Jackendoff, 2005</xref>; <xref ref-type="bibr" rid="B29">Clark, 2012</xref>; <xref ref-type="bibr" rid="B61">Hauser et al., 2014</xref>; <xref ref-type="bibr" rid="B7">Arbib, 2015</xref>). Thus, seminal work by <xref ref-type="bibr" rid="B146">Savage-Rumbaugh et al. (1986</xref>, <xref ref-type="bibr" rid="B149">2001</xref>; <xref ref-type="bibr" rid="B148">Savage-Rumbaugh and Rumbaugh, 1978</xref>) revealed that training chimpanzees on paired symbols designating items and actions (&#x201C;drink&#x201D; and &#x201C;liquids&#x201D; vs. &#x201C;give&#x201D; and &#x201C;solid foods&#x201D;) facilitated the learning of combinations of novel signs. In other words, the individuals more easily acquired pairs where action symbols correctly matched signs for types of foods, which implies a processing of signs in terms of their conceptual relations (<xref ref-type="bibr" rid="B39">Deacon, 1997</xref>, p. 86). More recently, <xref ref-type="bibr" rid="B92">Livingstone et al. (2014)</xref> trained rhesus monkeys on symbols representing distinct numbers of drops of liquid (implying a coding of magnitude). On tests involving combinations of these learned symbols, the individuals not only showed a capacity to process the <italic>relative</italic> values of signs within a context, but also transferred these subjective valuations to new symbols, suggesting a capacity to process combined signs in terms of novel relations. <xref ref-type="bibr" rid="B91">Livingstone et al. (2010</xref>, <xref ref-type="bibr" rid="B92">2014</xref>) also outlined that value coding of signs involves similar neural processes in humans and monkeys implicating dopamine neurons and interactions between the midbrain, the orbitofrontal cortex, and nucleus accumbens (for a critical review of other findings of this type, see <xref ref-type="bibr" rid="B116">N&#x00FA;&#x00F1;ez, 2017</xref>).</p>
<p>Finally, it was also believed that only humans have the ability to <italic>imitate</italic>, while apes <italic>emulate</italic> behaviors (e.g.,<xref ref-type="bibr" rid="B164">Tomasello, 1990</xref>, <xref ref-type="bibr" rid="B165">1996</xref>; <xref ref-type="bibr" rid="B63">Heyes, 1993</xref>; <xref ref-type="bibr" rid="B50">Fitch, 2010</xref>). Emulation has been characterized as entailing a learning of the effects of actions, rather than a copying of bodily motions. A limited capacity to imitate was thought to hinder the cultural transmission of communicative signs and tool use. Even so, some studies have shown that non-human primates can mimic the actions of their conspecifics and also learn to produce their calls and symbols (<xref ref-type="bibr" rid="B161">Sutton et al., 1973</xref>; <xref ref-type="bibr" rid="B54">Gardner and Gardner, 1985</xref>, <xref ref-type="bibr" rid="B55">1969</xref>; <xref ref-type="bibr" rid="B122">Pedersen, 2012</xref>). Recent research has made it clear that apes can selectively apply a range of social learning processes. This includes <italic>deferred imitation</italic> as well as the ability to follow eye-gaze and direct attention by eye-gaze and pointing (for a review, see <xref ref-type="bibr" rid="B171">Whiten et al., 2009</xref>). Other related claims to the effect that only humans have &#x201C;shared intentionality&#x201D; and an advanced &#x201C;theory of mind&#x201D; (e.g., <xref ref-type="bibr" rid="B131">Pinker and Jackendoff, 2005</xref>) have been questioned in studies of apes reared by humans (<xref ref-type="bibr" rid="B17">Bulloch et al., 2008</xref>; <xref ref-type="bibr" rid="B77">Leavens et al., 2009</xref>; see also contra <xref ref-type="bibr" rid="B124">Penn and Povinelli, 2007</xref>; <xref ref-type="bibr" rid="B18">Call and Tomasello, 2008</xref>).</p>
<p>In short, research in the last decades has revealed that, contrary to held assumptions, non-human primates possess mental abilities that serve to learn and process symbols. But the fact remains that monkeys and apes in the wild do not develop repertoires of symbolic signs of the type used in spoken language (e.g., <xref ref-type="bibr" rid="B129">Pika et al., 2005</xref>). To illustrate the kinds of signs that arise in non-human primates, one can consider the often cited case of vervet monkeys who use distinct signals to communicate the presence of different predators (<xref ref-type="bibr" rid="B153">Seyfarth et al., 1980</xref>; <xref ref-type="bibr" rid="B20">Cheney and Seyfarth, 1990</xref>; <xref ref-type="bibr" rid="B135">Price et al., 2015</xref>; for similar referent-specific signals in apes, see <xref ref-type="bibr" rid="B37">Crockford and Boesch, 2003</xref>). It has been argued that these signs are symbols based on their seeming arbitrariness (<xref ref-type="bibr" rid="B135">Price et al., 2015</xref>), though some critics reject this interpretation (<xref ref-type="bibr" rid="B59">Hauser et al., 2002</xref>; <xref ref-type="bibr" rid="B41">Deacon, 2012</xref>). In fact, vocal signs of apes and monkeys appear to be largely indexical in that they reflect reactions to referents in the signaling context (see also <xref ref-type="bibr" rid="B37">Crockford and Boesch, 2003</xref>; <xref ref-type="bibr" rid="B64">Hobaiter and Byrne, 2012</xref>; <xref ref-type="bibr" rid="B19">C&#x00E4;sar et al., 2013</xref>). Thus, research bears out that, while monkeys, apes, and humans share the cognitive abilities that are required to learn and use signs, only humans develop vast systems of symbols, and they do so primarily in a vocal medium. One implication of these results is that, even though cognitive abilities can be essential prerequisites in acquiring and manipulating symbols, some other capacity is needed to account for the emergence of these types of signs in vocal communication.</p>
</sec>
<sec><title>The Role of the Medium in the Rise of Symbolic Communication</title>
<sec><title>The Case Against Modality-Independent Accounts: &#x201C;Sign Languages&#x201D; and Storylines of the Gestural Origin of Language</title>
<p>Compared to cognitive skills, the capacity to articulate vocal patterns stands as an obvious human-specific trait. Yet in the literature on language origin, many researchers are guided by the belief that language emerged in a gestural medium. Several findings have motivated this view, which implies a conceptualization of &#x201C;language&#x201D; as an amodal function. One pivotal finding relates to the lack of voluntary control of vocalization in nonhuman primates.</p>
<p>In particular, studies by <xref ref-type="bibr" rid="B69">J&#x00FC;rgens (1976</xref>, <xref ref-type="bibr" rid="B70">1992</xref>) showed that the brains of monkeys and apes lack monosynaptic fibers linking the motor cortex to laryngeal-muscle motoneurons in the nucleus ambiguous. Such findings concur with the poor control over reactive vocalizations in these species (<xref ref-type="bibr" rid="B71">J&#x00FC;rgens, 2002</xref>, pp. 242&#x2013;245; <xref ref-type="bibr" rid="B155">Simonyan and Horwitz, 2011</xref>; <xref ref-type="bibr" rid="B154">Simonyan, 2014</xref>). The nervous systems of monkeys and apes do, however, present direct monosynaptic projections to motoneurons associated with the control of finger muscles, and to jaw- and lip-muscle motoneurons in the trigeminal and facial nuclei (<xref ref-type="bibr" rid="B71">J&#x00FC;rgens, 2002</xref>; <xref ref-type="bibr" rid="B143">Sasaki et al., 2004</xref>; <xref ref-type="bibr" rid="B80">Lemon, 2008</xref>). Compared to humans, though, there are fewer direct connections to tongue-muscle motoneurons in the hypoglossal nucleus (<xref ref-type="bibr" rid="B76">Kuypers, 1958</xref>; <xref ref-type="bibr" rid="B71">J&#x00FC;rgens, 2002</xref>), which accords with the paucity of oral segmentations or syllable-like patterns in the calls of non-human primates (<xref ref-type="bibr" rid="B81">Lieberman, 1968</xref>, <xref ref-type="bibr" rid="B85">2006a</xref>; <xref ref-type="bibr" rid="B90">Lieberman et al., 1992</xref>). Taken together, these observations may have led many to believe that, since apes and monkeys produce vocal signals as inflexible reactions, symbolic language instead evolved from controllable hand gestures (e.g., <xref ref-type="bibr" rid="B62">Hewes, 1996</xref>; <xref ref-type="bibr" rid="B31">Corballis, 2002</xref>, <xref ref-type="bibr" rid="B33">2009</xref>, <xref ref-type="bibr" rid="B35">2017</xref>; <xref ref-type="bibr" rid="B166">Tomasello and Zuberb&#x00FC;hler, 2002</xref>; <xref ref-type="bibr" rid="B4">Arbib, 2005</xref>, <xref ref-type="bibr" rid="B6">2012</xref>, <xref ref-type="bibr" rid="B7">2015</xref>; <xref ref-type="bibr" rid="B57">Gentilucci and Corballis, 2006</xref>; <xref ref-type="bibr" rid="B134">Pollick and de Waal, 2007</xref>; <xref ref-type="bibr" rid="B9">Arbib et al., 2008</xref>). This held belief, however, conflicts with the general observation that gestural signs of non-human primates do not function <italic>symbolically</italic> (and are mostly iconic or indexical). Thus, theories of the gestural origin of language are not supported by observations of extant species. Instead, the theories refer to <italic>indirect</italic> evidence seen to support storylines which essentially suggest that the last common ancestor (LCA) of the <italic>homo</italic> and <italic>pan</italic> (chimpanzee and bonobo) genera had, at some point, developed <italic>symbolic gestures</italic> from which spoken language evolved. This view, popularized by authors like Corballis and Arbib (as outlined below), has been criticized on fundamental grounds.</p>
<p>One objection bears on the theoretical significance given to &#x201C;sign languages.&#x201D; Proponents of gestural theories frequently refer to sign languages as illustrating the possibility of a gestural stage in language evolution (e.g., <xref ref-type="bibr" rid="B31">Corballis, 2002</xref>; <xref ref-type="bibr" rid="B6">Arbib, 2012</xref>, chapter 6). As such, this interpretation adheres to a view of language as deriving from an amodal faculty (e.g., <xref ref-type="bibr" rid="B21">Chomsky, 1965</xref>, <xref ref-type="bibr" rid="B23">2000</xref>, <xref ref-type="bibr" rid="B25">2006</xref>; and also <xref ref-type="bibr" rid="B130">Pinker, 1994</xref>; <xref ref-type="bibr" rid="B59">Hauser et al., 2002</xref>; <xref ref-type="bibr" rid="B58">Hauser, 2016</xref>). For example, gestural signs are seen to support the idea that &#x201C;the language faculty is not tied to specific sensory modalities&#x201D; (<xref ref-type="bibr" rid="B23">Chomsky, 2000</xref>, p. 121), and that &#x201C;discoveries about sign languages [&#x2026;] provide substantial evidence that externalization is modality independent&#x201D; (<xref ref-type="bibr" rid="B25">Chomsky, 2006</xref>, p. 22). However, such claims repeatedly disregard the fact that there is no known case where a community of normal hearers develops a sign language as a primary means of communication (<xref ref-type="bibr" rid="B45">Emmorey, 2005</xref>). Said differently, gestural signs as a primary system of communication generally appear where people share a pathology affecting the hearing modality &#x2013; which hardly supports the notion of an amodal language capacity. On the contrary, it suggests that, given the normal human ability to control <italic>both</italic> vocalization and hand gestures, symbolic communication links to a vocal-auditory modality with visual gestures having an accessory imagistic role (<xref ref-type="bibr" rid="B108">McNeill, 2005</xref>). It follows that an account of the rise of spoken language requires an explanation of how and why symbolic signs link to the vocal medium. But assuming the gestural origin of language leads instead to posit an evolutionary shift from gestural to vocal signs, which presents a conundrum for evolutionists.</p>
<p>To explain this briefly, one can refer to the theories of <xref ref-type="bibr" rid="B30">Corballis (1992</xref>, <xref ref-type="bibr" rid="B32">2003</xref>, <xref ref-type="bibr" rid="B33">2009</xref>, <xref ref-type="bibr" rid="B34">2010</xref>, <xref ref-type="bibr" rid="B35">2017</xref>; <xref ref-type="bibr" rid="B57">Gentilucci and Corballis, 2006</xref>), and <xref ref-type="bibr" rid="B4">Arbib (2005</xref>, <xref ref-type="bibr" rid="B5">2011</xref>, <xref ref-type="bibr" rid="B6">2012</xref>, <xref ref-type="bibr" rid="B7">2015</xref>, <xref ref-type="bibr" rid="B8">2017</xref>; <xref ref-type="bibr" rid="B9">Arbib et al., 2008</xref>). A critical claim of these proposals is that a left-lateralized control of hand gestures in area F5 of the monkey cortex evolved into a left-sided dominance for language, which the authors locate in Broca&#x2019;s area. Corballis and Arbib also refer to research showing that mirror neurons in F5 discharge when a monkey observes hand motions in others (see, e.g., <xref ref-type="bibr" rid="B137">Rizzolatti and Arbib, 1998</xref>; <xref ref-type="bibr" rid="B46">Ferrari et al., 2005</xref>). Both see in these responses a mechanism of action understanding, and conjecture that mirror neurons played a role in the shift from a gestural to a vocal-modality of communication. On how this shift occurred, it is speculated that, when the LCA descended from trees and developed bipedalism, there was a freeing of the hands allowing the development of expressive manual signs. At first, this led to putative iconic pantomimes that became conventionalized and symbolic (at a &#x201C;protosign&#x201D; stage), before &#x201C;protospeech&#x201D; developed. However, it is difficult to find in this narrative a working mechanism that converts gestures to vocal signs. For example, Arbib recently explained that pantomimes created an &#x201C;open-ended semantics&#x201D; which &#x201C;&#x2026;provides the adaptive pressure for increased control over the vocal apparatus&#x201D; (<xref ref-type="bibr" rid="B7">Arbib, 2015</xref>, pp. 612&#x2013;613; see also 2017, p. 144). By this account, the semantics of protosigns &#x201C;&#x2026;establishes the machinery that can then begin to develop protospeech, perhaps initially through the utility of creating novel sounds to match degrees of freedom of manual gestures (rising pitch could represent an upward movement of the hand)&#x201D; (2015, p. 613). Thus, the core explanation in this view is that <italic>semantics</italic> drove the evolution of vocalization, and the pairing of physiological parameters of hand control with those of articulatory, laryngeal, and respiratory systems of (proto-) speech. In this account, the example of iconic signs (of pitch increase and hand rising) hardly helps to understand how vast systems of <italic>symbolic</italic> signs emerged. Critics have also questioned whether any realistic model can be devised to &#x201C;translate&#x201D; hand motions into sequences of vocalized sounds (<xref ref-type="bibr" rid="B62">Hewes, 1996</xref>; <xref ref-type="bibr" rid="B100">MacNeilage, 2008/2010</xref>, pp. 287&#x2013;288).</p>
<p>In weighing the above scenario, one should note that it rests on the claim that a left-sided control of hand motions in the monkey cortex overlaps mirror neurons and Broca&#x2019;s area in humans. It has been reported, however, that activity in mirror neurons during the perception and production of hand motions is not left-lateralized (<xref ref-type="bibr" rid="B10">Aziz-Zadeh et al., 2006</xref>). More generally, one might question the <italic>a priori</italic> validity of theories where semantics is seen to drive the evolution of mechanisms of vocalization. Such views are not limited to gestural accounts. They extend to a variety of theories that focus on cognitive skills while overlooking modality-specific constraints and how they shape signals and signs. In this orientation, it is as if symbol systems generate from some amodal mental function.</p>
<p>For example, <xref ref-type="bibr" rid="B39">Deacon (1997</xref>, <xref ref-type="bibr" rid="B41">2012</xref>) suggests an account whereby signs created by apes (and children) first appear to have an indexical function. Then, when the signs are logically combined, they become more symbolic (<xref ref-type="bibr" rid="B39">Deacon, 1997</xref>, chapter 5). But again, apes in natural environments do not develop symbolic communication despite a capacity to combine given signs (as per the experiments of Savage-Rumbaugh et al., among others), so the question remains: how is it that symbolic signs emerge for humans and why is this linked primarily to the vocal medium? On these questions, <xref ref-type="bibr" rid="B39">Deacon (1997)</xref> basically offers a circular explanation: &#x201C;&#x2026;language must be viewed as its own prime mover. It is the author of a co-evolved complex of adaptations arrayed around a single core semiotic innovation&#x2026;&#x201D; (p. 44).</p>
<p>Aside from core semantics or semiotic functions, some theories also submit that the evolution of symbolic communication was driven by socio-cognitive functions. For example, the theory of &#x201C;interactional instinct&#x201D; suggests that, in language acquisition as in language evolution, children signal their intention to do something, and &#x201C;The intent becomes a symbol that the child expresses in an emotionally based interaction&#x2026;&#x201D; (<xref ref-type="bibr" rid="B79">Lee and Schumann, 2005</xref>, p. 7; <xref ref-type="bibr" rid="B78">Lee et al., 2009</xref>). In related proposals, constellations of mental skills including &#x201C;shared intentionality,&#x201D; &#x201C;perspective-taking,&#x201D; &#x201C;comprehension&#x201D; (etc.), along with &#x201C;thought processes&#x201D; (<xref ref-type="bibr" rid="B1">Ackermann et al., 2014</xref>), and &#x201C;purpose&#x201D; (<xref ref-type="bibr" rid="B40">Deacon, 2011</xref>) are evoked as driving factors (for an overview of these types of factors, see the proposal of <xref ref-type="bibr" rid="B132">Pleyer, 2017</xref>, and <xref ref-type="bibr" rid="B133">Pleyer and Winters, 2015</xref>). These accounts collectively imply what some have called &#x201C;mentalistic teleological&#x201D; principles (<xref ref-type="bibr" rid="B3">Allen, 1996/2009</xref>), which do not accord with accepted features of evolution. In particular, one feature holds that evolution reflects biological change in relation to physical aspects of the environment (see <xref ref-type="bibr" rid="B27">Christiansen and Chater, 2008</xref>). Accepting this, it is difficult to fathom how the evolution of anatomical structures of vocal communication would be driven by &#x201C;semantics,&#x201D; &#x201C;thoughts,&#x201D; &#x201C;intentions&#x201D; (etc.), without some basis in the sensory effects of physical signals and physiological processes of sign production. Indeed, if one <italic>defines</italic> symbols as entailing associations between concepts and signal elements, then the notion that symbolic language arose from amodal mental factors presents a contradiction in terms in that, in the absence of a modality of communication, there are no signals to which concepts can associate (but cf. the pronouncement that language has little to do with communication: <xref ref-type="bibr" rid="B24">Chomsky, 2002</xref>, pp. 76&#x2013;77; 2012, p. 11).</p>
</sec>
<sec><title>Modality-Dependent Accounts of the Rise of Symbolic Language</title>
<sec>
<title>Mimesis and procedural learning</title>
<p>Contrary to the above viewpoint, several authors have submitted that language first evolved in the vocal modality and that this medium imposes particular constraints on the formation of signs (e.g., <xref ref-type="bibr" rid="B82">Lieberman, 1984</xref>, <xref ref-type="bibr" rid="B83">2000</xref>, <xref ref-type="bibr" rid="B84">2003</xref>, <xref ref-type="bibr" rid="B87">2007</xref>, <xref ref-type="bibr" rid="B88">2016</xref>; <xref ref-type="bibr" rid="B99">MacNeilage, 1998</xref>, <xref ref-type="bibr" rid="B100">2008/2010</xref>; <xref ref-type="bibr" rid="B160">Studdert-Kennedy, 1998</xref>; <xref ref-type="bibr" rid="B101">MacNeilage and Davis, 2000</xref>, <xref ref-type="bibr" rid="B102">2015</xref>). This basically refutes claims that sign languages and spoken languages have similar structure reflecting an amodal mental capacity (e.g., <xref ref-type="bibr" rid="B12">Bellugi et al., 1989</xref>). Thus, <xref ref-type="bibr" rid="B100">MacNeilage (2008/2010</xref>, chapter 13) explained that vocal and gestural modalities shape signs quite differently: for instance, gestural signs are holistic and can involve simultaneous hand and body motions whereas, in spoken language, sounds are strictly sequential, and are constrained by articulatory and respiratory-phonatory systems. It is useful to note that such a viewpoint is echoed in recent studies of emerging sign language that explicitly focus on the link between signs and constraints on gestures. Most revealing is the work of Sandler et al. on <italic>Al-Sayyid Bedouin Sign Language</italic> (ABSL), which led to expressed reservations both on gestural accounts of language origin, and linguistic methods which have dominated research on sign languages.</p>
<p>Traditionally, the study of sign languages has relied on linguistic analyses and assumptions which focus on abstract phonological features, &#x201C;word&#x201D; categories and their syntax, but which entirely neglect modality-specific differences between gesture and sound production (<xref ref-type="bibr" rid="B100">MacNeilage, 2008/2010</xref>). Sandler et al. noted several difficulties in attempting to pigeon-hole gestures in terms of assumed linguistic features and units, and instead examined gestures as such (<xref ref-type="bibr" rid="B138">Sandler, 2012</xref>, <xref ref-type="bibr" rid="B139">2013</xref>, <xref ref-type="bibr" rid="B140">2017</xref>; <xref ref-type="bibr" rid="B141">Sandler et al., 2014</xref>). Their study of ABSL revealed a correspondence between developing motor aspects of gestures and the complexity of expressed concepts, which question the doctrine that spoken and gestural systems can derive from a common modality-independent function. As <xref ref-type="bibr" rid="B139">Sandler (2013</xref>, pp. 194&#x2013;195) remarked, in commenting Arbib&#x2019;s gesture theory of language origin &#x2013;</p>
<disp-quote>
<p>&#x2026;a different motor system controls language in each modality [gestural and vocal], and the relation between that system and the grammar is different as well. Considering the fundamental differences in motor systems, I am mindful of the reasoning of experts in the relation between motor control and cognition (<xref ref-type="bibr" rid="B42">Donald, 1991</xref>; <xref ref-type="bibr" rid="B100">MacNeilage, 2008/2010</xref>) who insist on the importance of the evolution of the supporting motor system in the evolution of language to the extent that &#x2018;mental representation cannot be fully understood without consideration of activities available to the body for building such representations&#x2026; [including the] dynamic characteristics of the production mechanism&#x2019; (<xref ref-type="bibr" rid="B38">Davis et al., 2002</xref>).</p>
</disp-quote>
<p>The findings also implied a reevaluation of linguistic assumptions on sign languages (<xref ref-type="bibr" rid="B138">Sandler, 2012</xref>, pp. 35&#x2013;36):</p>
<disp-quote>
<p>&#x2026;many believe that sign languages rapidly develop into a system that is very similar to that of spoken languages. Research charting the development of Nicaraguan Sign Language (e.g., <xref ref-type="bibr" rid="B151">Senghas, 1995</xref>; <xref ref-type="bibr" rid="B72">Kegl et al., 1999</xref>), which arose in a school beginning in the late 1970s, has convinced some linguists, such as Steven Pinker, that the language was &#x201C;created in one leap when the younger children were exposed to the pidgin signing of the older children&#x2026;&#x201D; (<xref ref-type="bibr" rid="B130">Pinker, 1994</xref>: 36). This is not surprising if, as Chomsky believes, &#x201C;&#x2026;language evolved, and is designed, primarily as an instrument of thought, with externalization a secondary process.&#x201D; (<xref ref-type="bibr" rid="B25">Chomsky, 2006</xref>: 22). [&#x2026;] Our work on ABSL, of which the present study is a part, suggests that those of us who contributed to this general picture may have overstated our case.</p>
</disp-quote>
<p>In sum, motor aspects of gesture and sound production shape symbol systems differently and storylines suggesting that vocal symbols developed from gestural signs bear intractable problems of gesture-to-sound conversion. But while such problems suggest a necessary link between symbolic language and vocal processes, not all modality-specific accounts deal with the rise of symbolic signs.</p>
<p>As an example, Lieberman&#x2019;s proposal focuses on how spoken language, viewed as a combinatorial system, links to the evolution of speech processes in conjunction with the basal ganglia and its function as a &#x201C;sequencing engine&#x201D; (e.g., <xref ref-type="bibr" rid="B84">Lieberman, 2003</xref>, <xref ref-type="bibr" rid="B86">2006b</xref>, <xref ref-type="bibr" rid="B88">2016</xref>). However, this proposal does not specifically address the issue of how vocal symbols emerged. MacNeilage, on the other hand, views symbols as the most fundamental factor of language evolution (<xref ref-type="bibr" rid="B100">MacNeilage, 2008/2010</xref>, p. 137). In his &#x201C;frame/content&#x201D; theory, the vocal modality provided a prototypal frame, the syllable, which is seen to originate in cyclical motions such as mastication (<xref ref-type="bibr" rid="B99">MacNeilage, 1998</xref>, but see <xref ref-type="bibr" rid="B16">Boucher, 2007</xref> who shows that cyclicity in speech articulation has little to do with mastication or chewing motions). MacNeilage argues that, for arbitrary symbols to emerge, &#x201C;&#x2026;hominids needed to evolve a capacity to invent expressive conventions&#x201D; (p. 99), though this did not arise from a &#x201C;higher-order word-making capacity&#x201D; (as in <xref ref-type="bibr" rid="B59">Hauser et al., 2002</xref>). Instead, the capacity to form &#x201C;words&#x201D; rests on mimesis and procedural learning, which refines actions and memory of actions. <xref ref-type="bibr" rid="B43">Donald (1997)</xref> described these processes as a capacity to &#x201C;&#x2026;Rehearse the action, observe its consequences, remember these, then alter the form of the original act, varying one or more parameters, dictated by the memory of the consequences of the previous action, or by an idealized image of the outcome&#x201D; (p. 142). MacNeilage, quoting Donald, submits that while human infants manifest procedural learning at the babbling stage, great apes do not, so that &#x201C;It would be no exaggeration to say that this capacity is uniquely human, and forms the background for the whole of human culture including language&#x201D; (Donald, p. 142). Yet studies of tool manufacture in chimpanzees challenge this claim.</p>
<p>For instance, in a recent longitudinal study, <xref ref-type="bibr" rid="B168">Vale et al. (2016</xref>, <xref ref-type="bibr" rid="B167">2017</xref>) observed that chimpanzees are not only successful in creating tools to retrieve rewards. They can also retain a procedure of tool manufacture for years and transfer this knowledge to new tasks, indicating an ability to acquire skills. Again, procedural learning, as other cognitive abilities, is required in symbol learning. However, it is not a sufficient factor in accounting for the human-specific development of symbolic language in a vocal medium. Nonetheless, the above proposals share the view that symbolic communication does not emerge from mental functions alone but essentially links to a capacity to produce patterns like syllables and babble.</p>
</sec>
<sec>
<title>&#x201C;Sound symbolism&#x201D;: Questions of efficiency and ease of learning of iconic signs</title>
<p>Another theory which also rests on mimesis suggests that symbolic language may originate from iconic vocal signs (or sound symbols) on the assumption that &#x201C;iconicity seems easier&#x201D; than making arbitrary associations (<xref ref-type="bibr" rid="B74">Kita, 1997</xref>; <xref ref-type="bibr" rid="B111">Monaghan et al., 2012</xref>; <xref ref-type="bibr" rid="B163">Thompson et al., 2012</xref>; <xref ref-type="bibr" rid="B66">Imai and Kita, 2014</xref>; <xref ref-type="bibr" rid="B152">Sereno, 2014</xref>; <xref ref-type="bibr" rid="B125">Perlman et al., 2015</xref>; <xref ref-type="bibr" rid="B106">Massaro and Perlman, 2017</xref>). The idea that symbols arose from a mimicking of objects and events partly draws from experiments by <xref ref-type="bibr" rid="B142">Sapir (1929)</xref> and <xref ref-type="bibr" rid="B75">K&#x00F6;hler (1947)</xref> on &#x201C;sound-shape&#x201D; pairings, where listeners judge perceived consonants and vowels as relating, for instance, to &#x201C;angular&#x201D; and &#x201C;rounded&#x201D; forms, or <xref ref-type="bibr" rid="B117">Ohala&#x2019;s (1994)</xref> &#x201C;frequency code&#x201D; where pitch is related to features like &#x201C;size&#x201D; and &#x201C;brilliance&#x201D; (e.g., <xref ref-type="bibr" rid="B107">Maurer et al., 2006</xref>; the frequency code also extends across species that use vocal pitch in signaling aggressive/passive intentions; <xref ref-type="bibr" rid="B113">Morton, 1994</xref>). The central assumption is that such iconic cues are vital to the ontogenesis and phylogenesis of language because they inherently facilitate the sound-meaning mappings and the displacement of signs (e.g., <xref ref-type="bibr" rid="B112">Monaghan et al., 2014</xref>). Experiments showing sound-form associations in infants and adult learners are often cited as supporting this view. For instance, a study by <xref ref-type="bibr" rid="B169">Walker et al. (2010)</xref> shows that pre-babbling infants of four months are able to associate pitch to features such as height and brightness (see also <xref ref-type="bibr" rid="B111">Monaghan et al., 2012</xref>). However, the experiments do not serve to demonstrate the facilitating effect of iconic sounds on language development, or the necessity of an iconic stage in developing arbitrary signs of oral language.</p>
<p>On these issues, attempts to relate iconic signs to spoken language face a logical problem. Iconic gestures or sounds offer highly restricted sets of signs compared to arbitrary symbols. Any restriction on the number of signs will inherently limit the diversity of form-referent associations and thus the <italic>efficiency</italic> of signs in communicating fine distinctions in meaning, leading some to see that &#x201C;While sound symbolism might be useful, it could impede word learning&#x2026;&#x201D; (<xref ref-type="bibr" rid="B66">Imai and Kita, 2014</xref>, p. 3; also <xref ref-type="bibr" rid="B109">Monaghan and Christiansen, 2006</xref>; <xref ref-type="bibr" rid="B111">Monaghan et al., 2012</xref>). This is evident in signs serving to <italic>name</italic> referents where sound symbols are quite limited (e.g., it is difficult to conceive how one could mimic open sets of lexemes such as &#x201C;proper nouns&#x201D;). On the idea that vocal iconic signs may nonetheless be easier to learn than arbitrary signs, computational models and experiments involving adult learners lead to the opposite conclusion (<xref ref-type="bibr" rid="B56">Gasser, 2004</xref>; <xref ref-type="bibr" rid="B110">Monaghan et al., 2011</xref>, <xref ref-type="bibr" rid="B111">2012</xref>, <xref ref-type="bibr" rid="B112">2014</xref>). For instance, in a series of experiments, <xref ref-type="bibr" rid="B111">Monaghan et al. (2012</xref>, <xref ref-type="bibr" rid="B112">2014</xref>) compared the learning of arbitrary and iconic (systematic) sign-meaning pairs taking into account given co-occurring &#x201C;contextual&#x201D; elements. In both neural network simulations and behavioral experiments, arbitrary form-meaning pairs were learned with fewer errors and more rapidly than iconic pairs. Still, such tests do not address the issue of how <italic>arbitrary</italic> signs arise in the vocal medium, and why this is specific to humans.</p>
</sec>
<sec>
<title>Segmented vocalization and the rise of symbolic signs: A demonstration</title>
<p>The aforementioned simulations and tests focus on the learning of sound-shape associations by infants and adults with respect to the perception of <italic>given</italic> signs provided by an experimenter. Such protocols do not address the issue of how signs emerge, and infants in the first months of life may not produce symbolic signs. However, it is well-established that maturational changes in the vocal apparatus coincide with the rise of such signs in child speech. On this development, comparisons of the vocal processes of human infants and apes are revealing of the mechanisms underlying the rise of vocal symbols. In particular, pre-babbling infants, like nonhuman primates, are obligate nasal breathers and produce sounds with nasal resonances (<xref ref-type="bibr" rid="B114">Negus, 1949</xref>; <xref ref-type="bibr" rid="B36">Crelin, 1987</xref>; <xref ref-type="bibr" rid="B162">Thom et al., 2006</xref>). Acoustically, nasalization dampens upper harmonics and formants, reducing the distinctiveness of sounds. Moreover, continuous nasal air-flow during vocalization implies that articulatory motions may not create salient features of oral segmentation (<xref ref-type="bibr" rid="B81">Lieberman, 1968</xref>, <xref ref-type="bibr" rid="B82">1984</xref>; <xref ref-type="bibr" rid="B89">Lieberman et al., 1972</xref>; <xref ref-type="bibr" rid="B162">Thom et al., 2006</xref>). For instance, producing multiple elements like stops [p,t,k,b,d,g], fricatives [f,s,&#x0283;,v,z,&#x01B7;], or any articulatory pattern of segmentation requires modulations of oral pressure which are difficult to achieve in a system where air flows through the nose. For this reason, early productions of infants largely appear as continuous nasalized cries and vocalizations which, as in the vocal sounds of non-human primates, are divided by breath interruptions and glottal closures (<xref ref-type="bibr" rid="B36">Crelin, 1987</xref>; <xref ref-type="bibr" rid="B95">Lynch et al., 1995</xref>). At three months, though, humans manifest a control of pitch contours, which can reflect a refinement of monosynaptic connections between the motor-cortex and motoneurons of laryngeal muscles (<xref ref-type="bibr" rid="B1">Ackermann et al., 2014</xref>). Some suggest that, at this stage, vocalizations become less reactive and iconic, and can involve a symbolic coding of pitch (<xref ref-type="bibr" rid="B118">Oller, 2000</xref>; <xref ref-type="bibr" rid="B119">Oller and Griebel, 2014</xref>). Subsequent supraglottal changes occur which are also human specific.</p>
<p>Of interest is the progressive decoupling of the nasopharynx indexed by the distancing of the epiglottis from the velum. In human newborns, as in many mammals, the epiglottis habitually overlaps the velum creating a sealed passage that allows nasal breathing while ingesting food and extracting breast milk (<xref ref-type="bibr" rid="B114">Negus, 1949</xref>; <xref ref-type="bibr" rid="B73">Kent, 1984</xref>; <xref ref-type="bibr" rid="B36">Crelin, 1987</xref>). It should be noted that the decoupling of the nasopharynx involves <italic>soft</italic> tissues <italic>&#x2013;</italic> hence the difficulty of dating this change from fossil records. Many works of comparative anatomy involving CT and MRI scans discuss this decoupling by reference to a &#x201C;descent of the larynx&#x201D; which is indexed by a lowering of the hyoid bone attached to the larynx, and related measures of vocal-tract length. Because this lowering is observed across species, some have concluded that laryngeal descent has little to do the rise of spoken language in humans (e.g., <xref ref-type="bibr" rid="B52">Fitch and Reby, 2001</xref>; <xref ref-type="bibr" rid="B50">Fitch, 2010</xref>, chapter 8). However, laryngeal descent is only accompanied by a <italic>permanent</italic> decoupling of the epiglottis and the velum in humans, beginning at about 6&#x2013;8 months (<xref ref-type="bibr" rid="B73">Kent, 1984</xref>; <xref ref-type="bibr" rid="B115">Nishimura et al., 2006</xref>; <xref ref-type="bibr" rid="B87">Lieberman, 2007</xref>). Following this decoupling, articulatory motions can create a variety of salient oral features in vocal signals, which constitutes a pivotal human-specific development. Though some contend that other species can produce a range of vowel-like resonances (<xref ref-type="bibr" rid="B48">Fitch, 2000</xref>, <xref ref-type="bibr" rid="B49">2005</xref>; <xref ref-type="bibr" rid="B52">Fitch and Reby, 2001</xref>), only humans segment these resonances using varying articulatory motions. This important change is accompanied by a general increase in rhythmic behavior (<xref ref-type="bibr" rid="B67">Iverson et al., 2007</xref>) leading to <italic>babble</italic> (<xref ref-type="bibr" rid="B93">Locke, 1995</xref>; <xref ref-type="bibr" rid="B118">Oller, 2000</xref>). Combining this morphological development with a capacity to modulate pitch contours confers the unique ability to manipulate both tonal and articulatory patterns. As noted, compared to other primates, humans develop direct monosynaptic projections to motoneurons of laryngeal muscles and also present a greater ratio of direct connections to tongue-muscle motoneurons which together support a control of orally modulated patterns of vocalization.</p>
<p>The symbolic potential of these patterns can be fathomed by considering that, in canonical babble, reduplicated syllables containing articulatory features such as [dada], [mamama], [nana] (etc.) become rapidly associated with caregivers, food, and other contextual referents. These early signs show that the <italic>arbitrariness</italic> underlying symbolic language can be <italic>inherent</italic> to the types of sounds that arise with orally segmented speech (a point also noted by <xref ref-type="bibr" rid="B31">Corballis, 2002</xref>), and that contextual information suffices to establish functional sound-meaning associations. In fact, the developmental literature does not suggest that iconic sounds or gestural mimics precede the rise of symbolic signs in canonical and variegated babble (<xref ref-type="bibr" rid="B93">Locke, 1995</xref>; <xref ref-type="bibr" rid="B118">Oller, 2000</xref>). On the other hand, it is generally acknowledged that the rise of vocal symbols accompanies a shift from pitch-varying to orally segmented vocalizations, though some symbol coding of pitch can precede this shift (<xref ref-type="bibr" rid="B120">Oller and Lynch, 1992</xref>; <xref ref-type="bibr" rid="B118">Oller, 2000</xref>).</p>
<p>Overall, the above developments suggest that the human capacity to produce orally segmented vocalizations presents a <italic>necessary</italic> &#x2013; though not sufficient <italic>&#x2013;</italic> factor in the rise of symbolic signs. In other words, cognitive and perceptual abilities are certainly required in forming sound-meaning associations, but these abilities are present at a basic level in non-human primates who do not develop symbolic signs like those of spoken language. Hence, the particular shift from pitch-varying to articulated vocalizations appears essential to the emergence of efficient vocal symbols in that, compared to pitch patterns, oral modulations offer a more diverse set of salient articulatory features by which to create fine distinctions in meaning.</p>
<p>Of course, a direct demonstration of this shift on the development of vocal symbols is not possible (i.e., one may not experimentally manipulate the human ability to articulate sounds). But one can artificially reduce acoustic features associated with articulatory motions by filtering signals so as to observe effects on the formation of symbolic associations. This guided the design of a straightforward demonstration using a task where listeners learn to associate pictures to unfamiliar speech sounds (in this case, two-syllable &#x201C;words&#x201D; spoken in Mandarin). In the task, sounds are presented with sets of possible referents (pictures of familiar objects) allowing the rise of associations through repeated trials. To evaluate the effect of a shift from intoned to orally articulated patterns on symbolic associations, filtered stimuli are presented bearing only pitch-varying intonations, then unfiltered stimuli with oral features are presented (for an example of the stimuli, see &#x201C;Materials and methods&#x201D; Figure <xref ref-type="fig" rid="F3">3</xref>). To support word-picture associations, two types of feedback are used representing two basic responses that can be obtained in a context of verbal learning. In the first type (henceforth <italic>feedback A</italic>), learners make an association and obtain an indication on whether or not the association is correct (&#x201C;yes/no&#x201D;). In the second type (<italic>feedback B</italic>), learners additionally receive information on what the sound designates (the correct picture is displayed). The first feedback condition emphasizes a process of inference, while the second favors rote learning. In both conditions, the prediction was that symbolic sound-meaning associations would form more rapidly for orally segmented sounds than for intoned patterns principally because of the greater efficiency of oral features in making distinctions in meaning.</p>
</sec>
</sec></sec>
</sec>
<sec><title>Results</title>
<p>Figure <xref ref-type="fig" rid="F1">1</xref> shows the effect of a change from intoned to orally modulated patterns on the rise of symbolic associations. Upon hearing sets of items containing features of oral articulation, arbitrary sound-meaning associations shift and form more rapidly and accurately than when the items are heard with only their tonal attributes, and this effect appears across feedback conditions. A repeated-measures ANOVA confirmed main effects of stimuli type [<italic>F</italic>(1,39) = 2.202; <italic>p</italic> &#x003C; 0.001; &#x03b7;<sub>p</sub><sup>2</sup> = 0.822; <italic>MSE</italic> = 0.012], and feedback ([<italic>F</italic>(1,39) = 3.581; <italic>p</italic> &#x003C; 0.001; &#x03b7;<sub>p</sub><sup>2</sup> = 0.896; <italic>MSE</italic> = 0.011]. Orally segmented stimuli yielded more correct associations than filtered stimuli, and <italic>feedback B</italic> led to more correct associations than <italic>feedback A</italic>. There was no significant interaction between the type of contexts and feedback conditions [<italic>F</italic>(1,39) = 8.740E-5; n.s.; &#x03b7;<sub>p</sub><sup>2</sup> = 0.000; <italic>MSE</italic> = 0.012].</p>
<fig id="F1" position="float">
<label>FIGURE 1</label>
<caption><p>Percent correct sound-picture association for different sets of lexemes presented with and without a filtering of oral features. Note that, when intoned items are presented with their oral features, symbolic associations rise more quickly and more accurately across feedback conditions.</p></caption>
<graphic xlink:href="fpsyg-09-02300-g001.tif"/>
</fig>
<p>As a further illustration, a separate comparison of sound-meaning associations was performed for sets of filtered items with different tones of Mandarin. Half of the contexts had initial flat tones while the other half contained variable rising and falling tones. One could surmise that symbolic associations could be facilitated for sets of items with varying tones compared to items that have only flat tones. However, as Figure <xref ref-type="fig" rid="F2">2</xref> illustrates, this effect did not occur. A repeated measures ANOVA showed no main effects of tonal patterns [<italic>F</italic>(1,39) = 0.049; n.s.; &#x03b7;<sub>p</sub><sup>2</sup> = 0.069; <italic>MSE</italic> = 0.017], but a main effect of feedback [<italic>F</italic>(1,39) = 2.175; <italic>p</italic> &#x003C; 0.001; &#x03b7;<sub>p</sub><sup>2</sup> = 0.757; <italic>MSE</italic> = 0.018] with <italic>feedback B</italic> leading to more correct associations than <italic>feedback A</italic>. Again, there was no significant interaction between the types of contexts and feedback [<italic>F</italic>(1,39) = 0.000; n.s.; &#x03b7;<sub>p</sub><sup>2</sup> = 0.001; <italic>MSE</italic> = 0.013]. A visual comparison with Figure <xref ref-type="fig" rid="F2">2</xref> suggests that, overall, intonational features are less efficient in supporting the formation of symbolic associations than oral features. It should also be weighed that many languages unlike Mandarin do not code tone at all in distinguishing lexical items, and thus, from a general perspective, intonation as such may not suffice in distinguishing large repertoires of lexemes.</p>
<fig id="F2" position="float">
<label>FIGURE 2</label>
<caption><p>Percent correct sound-picture association for different sets of filtered lexemes. Items with varying (rising and falling) tones did not lead to more symbolic associations than items with only flat tones.</p></caption>
<graphic xlink:href="fpsyg-09-02300-g002.tif"/>
</fig>
</sec>
<sec><title>Discussion</title>
<p>The preceding results demonstrate a seemingly self-evident effect: beyond the competence of the above learners and their experience with language, they were more prone to form arbitrary sound-meaning associations with oral patterns of speech than when speech contained only intonational patterns. Specifically, when shifting from items with tonal features to items that include features of articulation, symbolic associations were formed more rapidly and accurately. Of course, these observations and the above test are not meant to reflect evolution. Their purpose is to demonstrate the basic point that one may not account for the rise of symbolic signs in language without some reference to the types of signals that a modality of expression affords. Yet this is not a generally accepted principle.</p>
<p>As seen in the above review, the idea that symbolic language evolved from modality-independent cognitive abilities is widespread. One should also remark that the definitions of symbols that guide much of the work on language evolution refer to 19th century writers like Peirce and Saussure who did not consider how processes of modality can shape signals and signs. Following this tradition, storylines of language origin are largely oriented by the belief, popularized in linguistic theory, that symbolic language derives from a mental competence or faculty which has little to do with processes of expression like gestures and speech (<xref ref-type="bibr" rid="B144">Saussure, 1916/1966</xref>; <xref ref-type="bibr" rid="B21">Chomsky, 1965</xref>). From this standpoint, the rise of symbolic language would seem &#x201C;mysterious&#x201D; (<xref ref-type="bibr" rid="B61">Hauser et al., 2014</xref>), &#x201C;puzzling&#x201D; (<xref ref-type="bibr" rid="B2">Aitchison, 2000</xref>; <xref ref-type="bibr" rid="B15">Bouchard, 2015</xref>), and can lead to ask &#x201C;why only us&#x201D; (<xref ref-type="bibr" rid="B13">Berwick and Chomsky, 2015</xref>) or to speculations of evolutionary saltations (e.g., <xref ref-type="bibr" rid="B59">Hauser et al., 2002</xref>). However, linguistic analyses that focus on abstract categories and units overlook the structural effects of varying modalities of expression. On these effects, we noted that work on emerging sign and spoken language bears out that motor aspects of gestures and vocalization do shape sign systems very differently. As for the idea that symbols arise from a mental capacity, decades of research has shown that monkeys and apes have a basic competence to learn, process, and combine symbolic signs. Yet they do not develop the types of productive vocal symbols found in spoken language. One implication is that the human specificity of symbolic language may not be explained in terms of cognitive capacities alone. Other factors relating to the human ability to control vocal signals are needed to account for symbolic language and why it arises primarily in oral medium.</p>
<p>On these factors, several works of comparative physiology have identified human-specific changes that can underlie the rise of symbolic signs, and these reflect in ontogenesis in terms of a shift from pitch-varying patterns to orally articulated babble. As <xref ref-type="bibr" rid="B31">Corballis (2002)</xref>, pp. 187&#x2013;188) indicated, articulatory modulations of sounds generate numerous features in signals creating inherently arbitrary signs. These signs can be rapidly associated with co-occurring referents in the course of developing language. No stage of iconic gestures or &#x201C;sound-symbolism&#x201D; appears to precede this development (<xref ref-type="bibr" rid="B93">Locke, 1995</xref>; <xref ref-type="bibr" rid="B118">Oller, 2000</xref>). But nor does such a stage seem necessary. In fact, several studies have shown that infants, even pre-babblers, readily associate heard &#x201C;words&#x201D; with referents, and use these symbols to categorize objects or parts of objects (see <xref ref-type="bibr" rid="B53">Fulkerson and Waxman, 2007</xref>; <xref ref-type="bibr" rid="B47">Ferry et al., 2010</xref>, and for children of 12 months, <xref ref-type="bibr" rid="B96">MacKenzie et al., 2011</xref>). Interestingly, the reports also show that infants are less successful in forming symbolic associations when presented with sounds like intoned [mmm] or sounds that cannot be articulated, and instead attend to familiar speech (<xref ref-type="bibr" rid="B96">MacKenzie et al., 2011</xref>; <xref ref-type="bibr" rid="B104">Marno et al., 2015</xref>, <xref ref-type="bibr" rid="B105">2016</xref>; and for evidence that activity in language areas of the brain are organized in terms of speech sounds, see <xref ref-type="bibr" rid="B103">Magrassi et al., 2015</xref>). This ability to acquire symbols is not distinctly human, as we noted, but communication by way of orally articulated signs is. One can only speculate on what might have evolved had humans been limited to pitch-varying calls. From the above results, it can be surmised that pitch-controlled signals without features of oral articulation would restrict the rapid formation of efficient (accurate) symbols. On the phylogenesis of this capacity, certainly a pivotal factor is the decoupling of the nasopharynx which contributed to free the oral tract allowing an articulatory modulation of sounds (e.g., <xref ref-type="bibr" rid="B114">Negus, 1949</xref>; <xref ref-type="bibr" rid="B82">Lieberman, 1984</xref>, <xref ref-type="bibr" rid="B86">2006b</xref>; <xref ref-type="bibr" rid="B36">Crelin, 1987</xref>). Some see this as a consequence of bipedalism (originating 5&#x2013;7 million years ago) and that bipedalism further led to a decoupling of respiration and locomotion that supported an independent control of phonation (<xref ref-type="bibr" rid="B136">Provine, 2005</xref>; see also <xref ref-type="bibr" rid="B97">Maclarnon and Hewitt, 1999</xref>, <xref ref-type="bibr" rid="B98">2004</xref>). However, as we mentioned, bone markers in fossil records may not serve to index the separation of the nasopharynx, so dating this change as co-evolving with bipedalism appears problematic.</p>
</sec>
<sec id="s1" sec-type="materials|methods">
<title>Materials and Methods</title>
<sec><title>Participants</title>
<p>The participants were 40 speakers aged 20 to 41 years (mean of 25 years; 19 females) with no history of hearing problems or knowledge of any tone language. Prior to the task, all were screened for normal memory performance using a standard digit-span test (<xref ref-type="bibr" rid="B170">Wechsler, 1997</xref>).</p>
</sec>
<sec><title>Speech and Picture Stimuli</title>
<p>The original speech stimuli consisted of 24 two-syllable Mandarin lexical items (&#x201C;words&#x201D;) that matched 24 black-and-white pictures of objects (<xref ref-type="bibr" rid="B156">Snodgrass and Vanderwart, 1980</xref>). Half of these contexts had an initial flat tone and half a varying rising or falling tone (used to compare effects of coded pitch). The items were produced in isolation by a native female speaker of Mandarin and recorded using a headset microphone (AKG, model C-477-WR) and an external sound card (Shure, model X2u) set to a 16-bit resolution and 44.1 kHz sampling rate. The recorded items were stored as separate .wav files, and copies of these files were low-passed filtered at 350 Hz using an IIR function (<italic>Goldwave</italic>, v. 6.19) giving 24 filtered and 24 unfiltered versions of each of the lexemes. Finally, all of the stimuli were amplitude normalized to obtain similar dB intensities.</p>
<p>Figure <xref ref-type="fig" rid="F3">3</xref> provides an example of the filtered and unfiltered version of a spoken lexeme where one can see that only the lower harmonic is present in the filtered version, which gave natural-sounding pitch patterns with a vocalic quality. The distinctiveness of these contexts was evaluated by an external judge (not the authors) using a discrimination test. This was done by creating duplicates (<italic>AA</italic>) of the filtered items and placing another filtered test item (<italic>X</italic>) with similar tones before and following the pair, giving series of <italic>AAX</italic> and <italic>XAA</italic>. All tokens <italic>X</italic> were correctly discriminated indicating that the filtered stimuli retained distinguishable elements.</p>
<fig id="F3" position="float">
<label>FIGURE 3</label>
<caption><p>Example of filtered (left) and unfiltered (right) versions of a two-syllable lexeme in Mandarin. The filtering leaves the pitch (F0) of the lower harmonic, and the item is heard as vocalized tones without &#x201C;vowel&#x201D; or &#x201C;consonant&#x201D; features.</p></caption>
<graphic xlink:href="fpsyg-09-02300-g003.tif"/>
</fig>
</sec>
<sec><title>Test Design and Procedure</title>
<p>A repeated measures design was used with a counterbalancing of blocks of spoken lexemes on two type conditions (filtered vs. unfiltered), and two feedback conditions (<italic>A</italic> vs. <italic>B</italic>, described earlier), while taking into account a specific order of presentation (filtered items with <italic>feedback A</italic> were presented before filtered items with <italic>feedback B</italic>; then, unfiltered items with <italic>feedback A</italic> were presented before unfiltered items with <italic>feedback B</italic>). The latter presentation sequence basically aimed to capture effects of a shift from tonal to orally segmented patterns. The 24 lexemes were arranged into four lists of trials, each containing six repetitions of six different lexemes, giving 36 test trials per list. Each trial lexeme within the lists was matched to an array of four pictures with one correct lexeme-picture match and three random fillers. The counterbalance design implied that, across participants, all lists were presented an equal number of times in each condition.</p>
<p>In the experiment, a test trial began with a three-second display of four pictures, followed by a heard speech stimulus. The participant then had to select a likely speech-picture match by pressing a number on a keyboard (a forced-choice task). After the participant&#x2019;s key-press response, a feedback slide was displayed for two seconds. During the test, participants sat in a sound-treated room in front of a monitor and listened with headphones (Beyerdynamic, model DT 250) to the speech stimuli which were delivered via software (<italic>E-Prime 2</italic>). The sound stimuli had peak values of 71 dBA at the ears, as measured with a sound-level meter and headphone adapter (Digital Recordings, model DR-1). Practice runs were provided prior to the test, which proceeded without interruption. Statistical analyses of the responses used the procedures of <italic>SPSS</italic> (v. 17.0).</p>
</sec>
</sec>
<sec><title>Ethics Statement</title>
<p>This study was carried out in accordance with the recommendations of the &#x2018;Declaration of Helsinki&#x2019; with written informed consent from all subjects. The protocol was approved by the Comit&#x00E9; d&#x2019;&#x00C9;thique de la Recherche de la Facult&#x00E9; d&#x2019;Arts et Sciences de l&#x2019;Universit&#x00E9; de Montr&#x00E9;al.</p>
</sec>
<sec><title>Author Contributions</title>
<p>VB conceived the theoretical arguments and, with AG, designed the test. AR-B contributed to the data acquisition and procedure. All three authors contributed to the data analysis, interpretation, critical revision, and final approval of the paper.</p>
</sec>
<sec><title>Conflict of Interest Statement</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
</body>
<back>
<fn-group>
<fn fn-type="financial-disclosure">
<p><bold>Funding.</bold> The research was supported in part by a grant of the <italic>Fonds Qu&#x00E9;b&#x00E9;cois de la Recherche sur la Nature et la Technologie</italic> (FQRNT No. 175811).</p>
</fn>
</fn-group>
<ref-list>
<title>References</title>
<ref id="B1"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Ackermann</surname> <given-names>H.</given-names></name> <name><surname>Hage</surname> <given-names>S. R.</given-names></name> <name><surname>Ziegler</surname> <given-names>W.</given-names></name></person-group> (<year>2014</year>). <article-title>Brain mechanisms of acoustic communication in humans and nonhuman primates: an evolutionary perspective.</article-title> <source><italic>Behav. Brain Sci.</italic></source> <volume>37</volume> <fpage>529</fpage>&#x2013;<lpage>546</lpage>. <pub-id pub-id-type="doi">10.1017/S0140525X13003099</pub-id> <pub-id pub-id-type="pmid">24827156</pub-id></citation></ref>
<ref id="B2"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Aitchison</surname> <given-names>J.</given-names></name></person-group> (<year>2000</year>). <source><italic>The Seeds of Speech: Language Origin and Evolution.</italic></source> <publisher-loc>Cambridge</publisher-loc>: <publisher-name>Cambridge University Press</publisher-name>.</citation></ref>
<ref id="B3"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Allen</surname> <given-names>C.</given-names></name></person-group> (<year>1996/2009</year>). <article-title>&#x201C;Teleological notions in biology,&#x201D; in</article-title> <source><italic>The Stanford Encyclopedia of Philosophy</italic></source>, <role>ed.</role> <person-group person-group-type="editor"><name><surname>Zalta</surname> <given-names>E. N.</given-names></name></person-group> (<publisher-loc>Stanford CA</publisher-loc>: <publisher-name>Stanford University</publisher-name>).</citation></ref>
<ref id="B4"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Arbib</surname> <given-names>M. A.</given-names></name></person-group> (<year>2005</year>). <article-title>From monkey-like action recognition to human language: an evolutionary framework for neurolinguistics.</article-title> <source><italic>Behav. Brain Sci.</italic></source> <volume>28</volume> <fpage>105</fpage>&#x2013;<lpage>124</lpage>. <pub-id pub-id-type="doi">10.1017/S0140525X05000038</pub-id></citation></ref>
<ref id="B5"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Arbib</surname> <given-names>M. A.</given-names></name></person-group> (<year>2011</year>). <article-title>From mirror neurons to complex imitation in the evolution of language and tool use.</article-title> <source><italic>Anu. Rev. Anthropol.</italic></source> <volume>40</volume> <fpage>257</fpage>&#x2013;<lpage>273</lpage>. <pub-id pub-id-type="doi">10.1146/annurev-anthro-081309-145722</pub-id></citation></ref>
<ref id="B6"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Arbib</surname> <given-names>M. A.</given-names></name></person-group> (<year>2012</year>). <source><italic>How the Brain got Language: The Mirror System Hypothesis.</italic></source> <publisher-loc>Oxford</publisher-loc>: <publisher-name>Oxford University Press</publisher-name>. <pub-id pub-id-type="doi">10.1093/acprof:osobl/9780199896684.001.0001</pub-id></citation></ref>
<ref id="B7"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Arbib</surname> <given-names>M. A.</given-names></name></person-group> (<year>2015</year>). <article-title>&#x201C;Language evolution,&#x201D; in</article-title> <source><italic>The Handbook of Language Emergence</italic></source>, <role>eds</role> <person-group person-group-type="editor"><name><surname>MacWhinney</surname> <given-names>B.</given-names></name> <name><surname>O&#x2019;Grady</surname> <given-names>W.</given-names></name></person-group> (<publisher-loc>Hoboken, NJ</publisher-loc>: <publisher-name>John Wiley &#x0026; Sons</publisher-name>), <fpage>600</fpage>&#x2013;<lpage>623</lpage>.</citation></ref>
<ref id="B8"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Arbib</surname> <given-names>M. A.</given-names></name></person-group> (<year>2017</year>). <article-title>Toward the language-ready brain: biological evolution and primate comparisons.</article-title> <source><italic>Psychon. Bull. Rev.</italic></source> <volume>24</volume> <fpage>142</fpage>&#x2013;<lpage>150</lpage>. <pub-id pub-id-type="doi">10.3758/s13423-016-1098-2</pub-id> <pub-id pub-id-type="pmid">27368635</pub-id></citation></ref>
<ref id="B9"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Arbib</surname> <given-names>M. A.</given-names></name> <name><surname>Liebal</surname> <given-names>K.</given-names></name> <name><surname>Pika</surname> <given-names>S.</given-names></name></person-group> (<year>2008</year>). <article-title>Primate vocalization, gesture, and the evolution of human language.</article-title> <source><italic>Curr. Anthropol.</italic></source> <volume>49</volume> <fpage>1053</fpage>&#x2013;<lpage>1076</lpage>. <pub-id pub-id-type="doi">10.1086/593015</pub-id></citation></ref>
<ref id="B10"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Aziz-Zadeh</surname> <given-names>L.</given-names></name> <name><surname>Koski</surname> <given-names>L.</given-names></name> <name><surname>Zaidel</surname> <given-names>E.</given-names></name> <name><surname>Mazziotta</surname> <given-names>J.</given-names></name> <name><surname>Iacoboni</surname> <given-names>M.</given-names></name></person-group> (<year>2006</year>). <article-title>Lateralization of the human mirror neuron system.</article-title> <source><italic>J. Neurosci.</italic></source> <volume>26</volume> <fpage>2964</fpage>&#x2013;<lpage>2970</lpage>. <pub-id pub-id-type="doi">10.1523/jneurosci.2921-05.2006</pub-id></citation></ref>
<ref id="B11"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Belin</surname> <given-names>P.</given-names></name></person-group> (<year>2006</year>). <article-title>Voice processing in human and non-human primates.</article-title> <source><italic>Philos. Trans. R. Soc. B Biol. Sci.</italic></source> <volume>361</volume> <fpage>2091</fpage>&#x2013;<lpage>2107</lpage>. <pub-id pub-id-type="doi">10.1098/rstb.2006.1933</pub-id> <pub-id pub-id-type="pmid">17118926</pub-id></citation></ref>
<ref id="B12"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Bellugi</surname> <given-names>U.</given-names></name> <name><surname>Poizner</surname> <given-names>H.</given-names></name> <name><surname>Klima</surname> <given-names>E. S.</given-names></name></person-group> (<year>1989</year>). <article-title>Language, modality and the brain.</article-title> <source><italic>Trends Neurosci.</italic></source> <volume>12</volume> <fpage>380</fpage>&#x2013;<lpage>388</lpage>. <pub-id pub-id-type="doi">10.1016/0166-2236(89)90076-3</pub-id></citation></ref>
<ref id="B13"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Berwick</surname> <given-names>R. C.</given-names></name> <name><surname>Chomsky</surname> <given-names>N.</given-names></name></person-group> (<year>2015</year>). <source><italic>Why Only us: Language and Evolution.</italic></source> <publisher-loc>Cambridge MA</publisher-loc>: <publisher-name>MIT Press</publisher-name>.</citation></ref>
<ref id="B14"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Bolhuis</surname> <given-names>J. J.</given-names></name> <name><surname>Tattersal</surname> <given-names>I.</given-names></name> <name><surname>Chomsky</surname> <given-names>N.</given-names></name> <name><surname>Berwick</surname> <given-names>R. C.</given-names></name></person-group> (<year>2012</year>). <article-title>How could language have evolved?</article-title> <source><italic>PLoS Biol.</italic></source> <volume>12</volume>:<issue>e1001934</issue>. <pub-id pub-id-type="doi">10.1371/journal.pbio.1001934</pub-id> <pub-id pub-id-type="pmid">25157536</pub-id></citation></ref>
<ref id="B15"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Bouchard</surname> <given-names>D.</given-names></name></person-group> (<year>2015</year>). <article-title>Brain readiness and the nature of language.</article-title> <source><italic>Front. Psychol.</italic></source> <volume>6</volume>:<issue>1376</issue>. <pub-id pub-id-type="doi">10.3389/fpsyg.2015.01376</pub-id> <pub-id pub-id-type="pmid">26441751</pub-id></citation></ref>
<ref id="B16"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Boucher</surname> <given-names>V. J.</given-names></name></person-group> (<year>2007</year>). <article-title>&#x201C;Serial-order control and grouping in speech: findings for a frame/content theory,&#x201D; in</article-title> <source><italic>Proceedings of the XVIth International Congress of Phonetic Sciences</italic></source>, <role>eds</role> <person-group person-group-type="editor"><name><surname>Trouvain</surname> <given-names>J.</given-names></name> <name><surname>Barry</surname> <given-names>W. J.</given-names></name></person-group> (<publisher-loc>Saarbr&#x00FC;cken</publisher-loc>: <publisher-name>ICPhS</publisher-name>), <fpage>413</fpage>&#x2013;<lpage>416</lpage>.</citation></ref>
<ref id="B17"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Bulloch</surname> <given-names>M. J.</given-names></name> <name><surname>Boysen</surname> <given-names>S. T.</given-names></name> <name><surname>Furlong</surname> <given-names>E. E.</given-names></name></person-group> (<year>2008</year>). <article-title>Visual attention and its relation to knowledge states in chimpanzees, <italic>Pan troglodytes</italic>.</article-title> <source><italic>Anim. Behav.</italic></source> <volume>76</volume> <fpage>1147</fpage>&#x2013;<lpage>1155</lpage>. <pub-id pub-id-type="doi">10.1016/j.anbehav.2008.01.033</pub-id></citation></ref>
<ref id="B18"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Call</surname> <given-names>J.</given-names></name> <name><surname>Tomasello</surname> <given-names>M.</given-names></name></person-group> (<year>2008</year>). <article-title>Does the chimpanzee have a theory of mind? 30 years later.</article-title> <source><italic>Trends Cogn. Sci.</italic></source> <volume>12</volume> <fpage>187</fpage>&#x2013;<lpage>192</lpage>. <pub-id pub-id-type="doi">10.1016/j.tics.2008.02.010</pub-id> <pub-id pub-id-type="pmid">18424224</pub-id></citation></ref>
<ref id="B19"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>C&#x00E4;sar</surname> <given-names>C.</given-names></name> <name><surname>Zuberb&#x00FC;hler</surname> <given-names>K.</given-names></name> <name><surname>Young</surname> <given-names>R. J.</given-names></name> <name><surname>Byrne</surname> <given-names>R. W.</given-names></name></person-group> (<year>2013</year>). <article-title>Titi monkey call sequences vary with predator location and type.</article-title> <source><italic>Biol. Lett.</italic></source> <volume>9</volume> <fpage>1</fpage>&#x2013;<lpage>5</lpage>. <pub-id pub-id-type="doi">10.1098/rsbl.2013.0535</pub-id> <pub-id pub-id-type="pmid">24004492</pub-id></citation></ref>
<ref id="B20"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Cheney</surname> <given-names>D. L.</given-names></name> <name><surname>Seyfarth</surname> <given-names>R. M.</given-names></name></person-group> (<year>1990</year>). <source><italic>How Monkeys see the World.</italic></source> <publisher-loc>Chicago, IL</publisher-loc>: <publisher-name>University of Chicago Press</publisher-name>.</citation></ref>
<ref id="B21"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Chomsky</surname> <given-names>N.</given-names></name></person-group> (<year>1965</year>). <source><italic>Aspects of the theory of Syntax.</italic></source> <publisher-loc>Cambridge MA</publisher-loc>: <publisher-name>MIT Press</publisher-name>.</citation></ref>
<ref id="B22"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Chomsky</surname> <given-names>N.</given-names></name></person-group> (<year>1972</year>). <source><italic>Language and Mind.</italic></source> <publisher-loc>New York, NY</publisher-loc>: <publisher-name>Harcourt Brace Jovanovich</publisher-name>.</citation></ref>
<ref id="B23"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Chomsky</surname> <given-names>N.</given-names></name></person-group> (<year>2000</year>). <source><italic>New Horizons in the Study of Language and Mind.</italic></source> <publisher-loc>Cambridge</publisher-loc>: <publisher-name>Cambridge University Press</publisher-name>. <pub-id pub-id-type="doi">10.1017/CBO9780511811937</pub-id></citation></ref>
<ref id="B24"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Chomsky</surname> <given-names>N.</given-names></name></person-group> (<year>2002</year>). <source><italic>On Nature and Language.</italic></source> <publisher-loc>Cambridge</publisher-loc>: <publisher-name>Cambridge University Press</publisher-name>. <pub-id pub-id-type="doi">10.1017/CBO9780511613876</pub-id></citation></ref>
<ref id="B25"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Chomsky</surname> <given-names>N.</given-names></name></person-group> (<year>2006</year>). <source><italic>Language and mind.</italic></source> <publisher-loc>Cambridge</publisher-loc>: <publisher-name>Cambridge University Press</publisher-name>. <pub-id pub-id-type="doi">10.1017/CBO9780511791222</pub-id></citation></ref>
<ref id="B26"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Chomsky</surname> <given-names>N.</given-names></name></person-group> (<year>2012</year>). <source><italic>The Science of Language. Interviews with James McGilvray.</italic></source> <publisher-loc>Cambridge</publisher-loc>: <publisher-name>Cambridge University Press</publisher-name>. <pub-id pub-id-type="doi">10.1017/CBO9781139061018</pub-id></citation></ref>
<ref id="B27"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Christiansen</surname> <given-names>M. H.</given-names></name> <name><surname>Chater</surname> <given-names>N.</given-names></name></person-group> (<year>2008</year>). <article-title>Language as shaped by the brain.</article-title> <source><italic>Behav. Brain Sci.</italic></source> <volume>31</volume> <fpage>489</fpage>&#x2013;<lpage>558</lpage>. <pub-id pub-id-type="doi">10.1017/S0140525X08004998</pub-id> <pub-id pub-id-type="pmid">18826669</pub-id></citation></ref>
<ref id="B28"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Christiansen</surname> <given-names>M. H.</given-names></name> <name><surname>Kirby</surname> <given-names>S.</given-names></name></person-group> (<year>2003</year>). <article-title>Language evolution: consensus and controversies.</article-title> <source><italic>Trends Cogn. Sci.</italic></source> <volume>7</volume> <fpage>300</fpage>&#x2013;<lpage>307</lpage>. <pub-id pub-id-type="doi">10.1016/s1364-6613(03)00136-0</pub-id></citation></ref>
<ref id="B29"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Clark</surname> <given-names>B.</given-names></name></person-group> (<year>2012</year>). <article-title>Syntactic theory and the evolution of syntax.</article-title> <source><italic>Biolinguistics</italic></source> <volume>7</volume> <fpage>169</fpage>&#x2013;<lpage>197</lpage>.</citation></ref>
<ref id="B30"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Corballis</surname> <given-names>M. C.</given-names></name></person-group> (<year>1992</year>). <article-title>On the evolution of language and generativity.</article-title> <source><italic>Cognition</italic></source> <volume>44</volume> <fpage>197</fpage>&#x2013;<lpage>226</lpage>. <pub-id pub-id-type="doi">10.1016/0010-0277(92)90001-X</pub-id></citation></ref>
<ref id="B31"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Corballis</surname> <given-names>M. C.</given-names></name></person-group> (<year>2002</year>). <source><italic>From Hand to Mouth: The Origins of Language.</italic></source> <publisher-loc>Princeton NJ</publisher-loc>: <publisher-name>Princeton University Press</publisher-name>.</citation></ref>
<ref id="B32"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Corballis</surname> <given-names>M. C.</given-names></name></person-group> (<year>2003</year>). <article-title>From mouth to hand: gesture, speech, and the evolution of right-handedness.</article-title> <source><italic>Behav. Brain Sci.</italic></source> <volume>26</volume> <fpage>199</fpage>&#x2013;<lpage>208</lpage>; <comment>discussion</comment> <fpage>208</fpage>&#x2013;<lpage>160</lpage>. <pub-id pub-id-type="doi">10.1017/S0140525X03000062</pub-id> <pub-id pub-id-type="pmid">14621511</pub-id></citation></ref>
<ref id="B33"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Corballis</surname> <given-names>M. C.</given-names></name></person-group> (<year>2009</year>). <article-title>The evolution of language.</article-title> <source><italic>Ann. N. Y. Acad. Sci.</italic></source> <volume>1156</volume> <fpage>19</fpage>&#x2013;<lpage>43</lpage>. <pub-id pub-id-type="doi">10.1111/j.1749-6632.2009.04423.x</pub-id> <pub-id pub-id-type="pmid">19338501</pub-id></citation></ref>
<ref id="B34"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Corballis</surname> <given-names>M. C.</given-names></name></person-group> (<year>2010</year>). <article-title>Mirror neurons and the evolution of language.</article-title> <source><italic>Brain Lang.</italic></source> <volume>112</volume> <fpage>25</fpage>&#x2013;<lpage>35</lpage>. <pub-id pub-id-type="doi">10.1016/j.bandl.2009.02.002</pub-id> <pub-id pub-id-type="pmid">19342089</pub-id></citation></ref>
<ref id="B35"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Corballis</surname> <given-names>M. C.</given-names></name></person-group> (<year>2017</year>). <source><italic>The Truth about Language: What it is and Where it Came from.</italic></source> <publisher-loc>Chicago, IL</publisher-loc>: <publisher-name>University of Chicago Press</publisher-name>. <pub-id pub-id-type="doi">10.7208/chicago/9780226287225.001.0001</pub-id></citation></ref>
<ref id="B36"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Crelin</surname> <given-names>E. S.</given-names></name></person-group> (<year>1987</year>). <source><italic>The Human Vocal Tract: Anatomy, Function, Development and Evolution.</italic></source> <publisher-loc>New York, NY</publisher-loc>: <publisher-name>Vantage Press</publisher-name>.</citation></ref>
<ref id="B37"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Crockford</surname> <given-names>C.</given-names></name> <name><surname>Boesch</surname> <given-names>C.</given-names></name></person-group> (<year>2003</year>). <article-title>Context-specific calls in wild chimpanzees, <italic>Pan troglodytes</italic> verus: analysis of barks.</article-title> <source><italic>Anim. Behav.</italic></source> <volume>66</volume> <fpage>115</fpage>&#x2013;<lpage>125</lpage>. <pub-id pub-id-type="doi">10.1006/anbe.2003.2166</pub-id></citation></ref>
<ref id="B38"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Davis</surname> <given-names>B. L.</given-names></name> <name><surname>MacNeilage</surname> <given-names>P. F.</given-names></name> <name><surname>Matyear</surname> <given-names>C. L.</given-names></name></person-group> (<year>2002</year>). <article-title>Acquisition of serial complexity in speech production: a comparison of phonetic and phonological approaches to first word production.</article-title> <source><italic>Phonetica</italic></source> <volume>59</volume> <fpage>75</fpage>&#x2013;<lpage>107</lpage>. <pub-id pub-id-type="doi">10.1159/000066065</pub-id> <pub-id pub-id-type="pmid">12232462</pub-id></citation></ref>
<ref id="B39"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Deacon</surname> <given-names>T. W.</given-names></name></person-group> (<year>1997</year>). <source><italic>The Symbolic Species.</italic></source> <publisher-loc>New York, NY</publisher-loc>: <publisher-name>W.W. Norton</publisher-name>.</citation></ref>
<ref id="B40"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Deacon</surname> <given-names>T. W.</given-names></name></person-group> (<year>2011</year>). <source><italic>Incomplete Nature: How mind Emerged from Matter.</italic></source> <publisher-loc>New York, NY</publisher-loc>: <publisher-name>W.W. Norton</publisher-name>.</citation></ref>
<ref id="B41"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Deacon</surname> <given-names>T. W.</given-names></name></person-group> (<year>2012</year>). <article-title>&#x201C;Beyond the symbolic species,&#x201D; in</article-title> <source><italic>The Symbolic Species Evolved</italic></source>, <role>eds</role> <person-group person-group-type="editor"><name><surname>Schilhab</surname> <given-names>T.</given-names></name> <name><surname>Stjernfelt</surname> <given-names>F.</given-names></name> <name><surname>Deacon</surname> <given-names>T. W.</given-names></name></person-group> (<publisher-loc>Dordrecht</publisher-loc>: <publisher-name>Springer</publisher-name>), <fpage>9</fpage>&#x2013;<lpage>38</lpage>. <pub-id pub-id-type="doi">10.1007/978-94-007-2336-8_2</pub-id></citation></ref>
<ref id="B42"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Donald</surname> <given-names>M.</given-names></name></person-group> (<year>1991</year>). <source><italic>Origins of the Modern Mind: Three Stages in the Evolution of Culture and Cognition.</italic></source> <publisher-loc>Cambridge, MA</publisher-loc>: <publisher-name>Harvard University Press</publisher-name>.</citation></ref>
<ref id="B43"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Donald</surname> <given-names>M.</given-names></name></person-group> (<year>1997</year>). <article-title>&#x201C;Preconditions for the evolution of protolanguages,&#x201D; in</article-title> <source><italic>The Descent of mind: Psychological Perspectives on Hominid Evolution</italic></source>, <role>eds</role> <person-group person-group-type="editor"><name><surname>Corballis</surname> <given-names>M. C.</given-names></name> <name><surname>Lea</surname> <given-names>S. E. G.</given-names></name></person-group> (<publisher-loc>Oxford</publisher-loc>: <publisher-name>Oxford University Press</publisher-name>), <fpage>138</fpage>&#x2013;<lpage>154</lpage>.</citation></ref>
<ref id="B44"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Eichenbaum</surname> <given-names>H.</given-names></name> <name><surname>Cohen</surname> <given-names>N. J.</given-names></name></person-group> (<year>2001</year>). <source><italic>From Conditioning to Conscious Recollection: Memory Systems of the Brain.</italic></source> <publisher-loc>New York, NY</publisher-loc>: <publisher-name>Oxford University Press</publisher-name>.</citation></ref>
<ref id="B45"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Emmorey</surname> <given-names>K.</given-names></name></person-group> (<year>2005</year>). <article-title>Sign languages are problematic for a gestural origins theory of language evolution.</article-title> <source><italic>Behav. Brain Sci.</italic></source> <volume>28</volume> <fpage>130</fpage>&#x2013;<lpage>131</lpage>. <pub-id pub-id-type="doi">10.1017/S0140525X05270036</pub-id></citation></ref>
<ref id="B46"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Ferrari</surname> <given-names>P. F.</given-names></name> <name><surname>Rozzi</surname> <given-names>S.</given-names></name> <name><surname>Fogassi</surname> <given-names>L.</given-names></name></person-group> (<year>2005</year>). <article-title>Mirror neurons responding to observation of actions made with tools in monkey ventral premotor cortex.</article-title> <source><italic>J. Cogn. Neurosci.</italic></source> <volume>17</volume> <fpage>212</fpage>&#x2013;<lpage>226</lpage>. <pub-id pub-id-type="doi">10.1162/0898929053124910</pub-id> <pub-id pub-id-type="pmid">15811234</pub-id></citation></ref>
<ref id="B47"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Ferry</surname> <given-names>A. L.</given-names></name> <name><surname>Hespos</surname> <given-names>S. J.</given-names></name> <name><surname>Waxman</surname> <given-names>S. R.</given-names></name></person-group> (<year>2010</year>). <article-title>Categorization in 3-and 4-month-old infants: an advantage of words over tones.</article-title> <source><italic>Child Dev.</italic></source> <volume>81</volume> <fpage>472</fpage>&#x2013;<lpage>479</lpage>. <pub-id pub-id-type="doi">10.1111/j.1467-8624.2009.01408.x</pub-id> <pub-id pub-id-type="pmid">20438453</pub-id></citation></ref>
<ref id="B48"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Fitch</surname> <given-names>W. T.</given-names></name></person-group> (<year>2000</year>). <article-title>The evolution of speech: a comparative review.</article-title> <source><italic>Trends Cogn. Sci.</italic></source> <volume>4</volume> <fpage>258</fpage>&#x2013;<lpage>267</lpage>. <pub-id pub-id-type="doi">10.1016/S1364-6613(00)01494-7</pub-id></citation></ref>
<ref id="B49"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Fitch</surname> <given-names>W. T.</given-names></name></person-group> (<year>2005</year>). <article-title>&#x201C;Production of vocalizations in mammals,&#x201D; in</article-title> <source><italic>Encyclopedia of Language and Linguistics</italic></source>, <role>ed.</role> <person-group person-group-type="editor"><name><surname>Brown</surname> <given-names>K.</given-names></name></person-group> (<publisher-loc>New York, NY</publisher-loc>: <publisher-name>Elsevier</publisher-name>), <fpage>115</fpage>&#x2013;<lpage>121</lpage>.</citation></ref>
<ref id="B50"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Fitch</surname> <given-names>W. T.</given-names></name></person-group> (<year>2010</year>). <source><italic>The Evolution of Language.</italic></source> <publisher-loc>Cambridge</publisher-loc>: <publisher-name>Cambridge University Press</publisher-name>. <pub-id pub-id-type="doi">10.1017/CBO9780511817779</pub-id></citation></ref>
<ref id="B51"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Fitch</surname> <given-names>W. T.</given-names></name> <name><surname>Hauser</surname> <given-names>M. D.</given-names></name> <name><surname>Chomsky</surname> <given-names>N.</given-names></name></person-group> (<year>2005</year>). <article-title>The evolution of the language faculty: clarifications and implications.</article-title> <source><italic>Cognition</italic></source> <volume>97</volume> <fpage>179</fpage>&#x2013;<lpage>210</lpage>; <comment>discussion</comment> <fpage>211</fpage>&#x2013;<lpage>125</lpage>. <pub-id pub-id-type="doi">10.1016/j.cognition.2005.02.005</pub-id> <pub-id pub-id-type="pmid">16112662</pub-id></citation></ref>
<ref id="B52"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Fitch</surname> <given-names>W. T.</given-names></name> <name><surname>Reby</surname> <given-names>D.</given-names></name></person-group> (<year>2001</year>). <article-title>The descended larynx is not uniquely human.</article-title> <source><italic>Proc. R. Soc. Lond. B Biol. Sci.</italic></source> <volume>268</volume> <fpage>1669</fpage>&#x2013;<lpage>1675</lpage>. <pub-id pub-id-type="doi">10.1098/rspb.2001.1704</pub-id> <pub-id pub-id-type="pmid">11506679</pub-id></citation></ref>
<ref id="B53"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Fulkerson</surname> <given-names>A. L.</given-names></name> <name><surname>Waxman</surname> <given-names>S. R.</given-names></name></person-group> (<year>2007</year>). <article-title>Words (but not tones) facilitate object categorization: evidence from 6-and 12-month-olds.</article-title> <source><italic>Cognition</italic></source> <volume>105</volume> <fpage>218</fpage>&#x2013;<lpage>228</lpage>. <pub-id pub-id-type="doi">10.1016/j.cognition.2006.09.005</pub-id> <pub-id pub-id-type="pmid">17064677</pub-id></citation></ref>
<ref id="B54"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Gardner</surname> <given-names>B. T.</given-names></name> <name><surname>Gardner</surname> <given-names>R. A.</given-names></name></person-group> (<year>1985</year>). <article-title>Signs of Intelligence in cross-fostered chimpanzees.</article-title> <source><italic>Philos. Trans. R. Soc. Lond. B Biol. Sci.</italic></source> <volume>308</volume> <fpage>159</fpage>&#x2013;<lpage>176</lpage>. <pub-id pub-id-type="doi">10.1098/rstb.1985.0017</pub-id></citation></ref>
<ref id="B55"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Gardner</surname> <given-names>R. A.</given-names></name> <name><surname>Gardner</surname> <given-names>B. T.</given-names></name></person-group> (<year>1969</year>). <article-title>Teaching sign language to a chimpanzee.</article-title> <source><italic>Science</italic></source> <volume>165</volume> <fpage>664</fpage>&#x2013;<lpage>672</lpage>. <pub-id pub-id-type="doi">10.1126/science.165.3894.664</pub-id></citation></ref>
<ref id="B56"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Gasser</surname> <given-names>M.</given-names></name></person-group> (<year>2004</year>). <article-title>&#x201C;The origins of arbitrariness in language,&#x201D; in</article-title> <source><italic>Proceedings of the 26th Annual Meeting of the Cognitive Science Society</italic></source>, <role>eds</role> <person-group person-group-type="editor"><name><surname>Forbus</surname> <given-names>K. D.</given-names></name> <name><surname>Gentner</surname> <given-names>D.</given-names></name> <name><surname>Regier</surname> <given-names>T.</given-names></name></person-group> (<publisher-loc>Mahwah, NJ</publisher-loc>: <publisher-name>Lawrence Erlbaum</publisher-name>),<fpage>434</fpage>&#x2013;<lpage>439</lpage>.</citation></ref>
<ref id="B57"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Gentilucci</surname> <given-names>M.</given-names></name> <name><surname>Corballis</surname> <given-names>M. C.</given-names></name></person-group> (<year>2006</year>). <article-title>From manual gesture to speech: a gradual transition.</article-title> <source><italic>Neurosci. Biobehav. Rev.</italic></source> <volume>30</volume> <fpage>949</fpage>&#x2013;<lpage>960</lpage>. <pub-id pub-id-type="doi">10.1016/j.neubiorev.2006.02.004</pub-id> <pub-id pub-id-type="pmid">16620983</pub-id></citation></ref>
<ref id="B58"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Hauser</surname> <given-names>M. D.</given-names></name></person-group> (<year>2016</year>). <article-title>Challenges to the what, when, and why?</article-title> <source><italic>Biolinguistics</italic></source> <volume>10</volume> <fpage>1</fpage>&#x2013;<lpage>5</lpage>.</citation></ref>
<ref id="B59"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Hauser</surname> <given-names>M. D.</given-names></name> <name><surname>Chomsky</surname> <given-names>N.</given-names></name> <name><surname>Fitch</surname> <given-names>W. T.</given-names></name></person-group> (<year>2002</year>). <article-title>The faculty of language: what is it, who has it, and how did it evolve?</article-title> <source><italic>Science</italic></source> <volume>298</volume> <fpage>1569</fpage>&#x2013;<lpage>1579</lpage>.</citation></ref>
<ref id="B60"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Hauser</surname> <given-names>M. D.</given-names></name> <name><surname>Newport</surname> <given-names>E. L.</given-names></name> <name><surname>Aslin</surname> <given-names>R. N.</given-names></name></person-group> (<year>2001</year>). <article-title>Segmentation of the speech stream in a nonhuman primate: statistical learning in cotton-top tamarins.</article-title> <source><italic>Cognition</italic></source> <volume>78</volume> <fpage>B53</fpage>&#x2013;<lpage>B64</lpage>. <pub-id pub-id-type="doi">10.1016/S0010-0277(00)00132-3</pub-id></citation></ref>
<ref id="B61"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Hauser</surname> <given-names>M. D.</given-names></name> <name><surname>Yang</surname> <given-names>C.</given-names></name> <name><surname>Berwick</surname> <given-names>R. C.</given-names></name> <name><surname>Tattersall</surname> <given-names>I.</given-names></name> <name><surname>Ryan</surname> <given-names>M.</given-names></name> <name><surname>Watumull</surname> <given-names>J.</given-names></name><etal/></person-group> (<year>2014</year>). <article-title>The mystery of language evolution.</article-title> <source><italic>Front. Psychol.</italic></source> <volume>5</volume>:<issue>401</issue>. <pub-id pub-id-type="doi">10.3389/fpsyg.2014.00401</pub-id> <pub-id pub-id-type="pmid">24847300</pub-id></citation></ref>
<ref id="B62"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Hewes</surname> <given-names>G. W.</given-names></name></person-group> (<year>1996</year>). <article-title>&#x201C;A history of the study of language origins and the gestural primacy hypothesis,&#x201D; in</article-title> <source><italic>Handbook of Human Symbolic Evolution</italic></source>, <role>eds</role> <person-group person-group-type="editor"><name><surname>Lock</surname> <given-names>A.</given-names></name> <name><surname>Peters</surname> <given-names>C.</given-names></name></person-group> (<publisher-loc>Oxford</publisher-loc>: <publisher-name>Oxford University Press</publisher-name>), <fpage>571</fpage>&#x2013;<lpage>595</lpage>.</citation></ref>
<ref id="B63"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Heyes</surname> <given-names>C. M.</given-names></name></person-group> (<year>1993</year>). <article-title>Imitation, culture and cognition.</article-title> <source><italic>Anim. Behav.</italic></source> <volume>46</volume> <fpage>999</fpage>&#x2013;<lpage>1010</lpage>. <pub-id pub-id-type="doi">10.1006/anbe.1993.1281</pub-id></citation></ref>
<ref id="B64"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Hobaiter</surname> <given-names>C. L.</given-names></name> <name><surname>Byrne</surname> <given-names>R. W.</given-names></name></person-group> (<year>2012</year>). <article-title>&#x201C;Gesture use in consortship: wild chimpanzees&#x2019; use of gesture for an &#x2018;evolutionarily urgent&#x2019; purpose,&#x201D; in</article-title> <source><italic>Developments in Primate Gesture Research</italic></source>, <role>eds</role> <person-group person-group-type="editor"><name><surname>Pika</surname> <given-names>S.</given-names></name> <name><surname>Liebal</surname> <given-names>K.</given-names></name></person-group> (<publisher-loc>Amsterdam</publisher-loc>: <publisher-name>John Benjamins</publisher-name>), <fpage>129</fpage>&#x2013;<lpage>146</lpage>. <pub-id pub-id-type="doi">10.1075/gs.6.08hob</pub-id></citation></ref>
<ref id="B65"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Hockett</surname> <given-names>C. F.</given-names></name></person-group> (<year>1960</year>). <article-title>The origin of speech.</article-title> <source><italic>Sci. Am.</italic></source> <volume>203</volume> <fpage>88</fpage>&#x2013;<lpage>111</lpage>. <pub-id pub-id-type="doi">10.1038/scientificamerican0960-88</pub-id></citation></ref>
<ref id="B66"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Imai</surname> <given-names>M.</given-names></name> <name><surname>Kita</surname> <given-names>S.</given-names></name></person-group> (<year>2014</year>). <article-title>The sound symbolism bootstrapping hypothesis for language acquisition and language evolution.</article-title> <source><italic>Philos. Trans. R. Soc. B Biol. Sci.</italic></source> <volume>369</volume> <fpage>1</fpage>&#x2013;<lpage>13</lpage>. <pub-id pub-id-type="doi">10.1098/rstb.2013.0298</pub-id> <pub-id pub-id-type="pmid">25092666</pub-id></citation></ref>
<ref id="B67"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Iverson</surname> <given-names>J. M.</given-names></name> <name><surname>Hall</surname> <given-names>A. J.</given-names></name> <name><surname>Nickel</surname> <given-names>L.</given-names></name> <name><surname>Wozniak</surname> <given-names>R. H.</given-names></name></person-group> (<year>2007</year>). <article-title>The relationship between reduplicated babble onset and laterality biases in infant rhythmic arm movements.</article-title> <source><italic>Brain Lang.</italic></source> <volume>101</volume> <fpage>198</fpage>&#x2013;<lpage>207</lpage>. <pub-id pub-id-type="doi">10.1016/j.bandl.2006.11.004</pub-id> <pub-id pub-id-type="pmid">17196644</pub-id></citation></ref>
<ref id="B68"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Jackendoff</surname> <given-names>R.</given-names></name></person-group> (<year>2002</year>). <source><italic>Foundations of Language: Brain, Meaning, Grammar, Evolution.</italic></source> <publisher-loc>New York, NY</publisher-loc>: <publisher-name>Oxford University Press</publisher-name>. <pub-id pub-id-type="doi">10.1093/acprof:oso/9780198270126.001.0001</pub-id></citation></ref>
<ref id="B69"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>J&#x00FC;rgens</surname> <given-names>U.</given-names></name></person-group> (<year>1976</year>). <article-title>Projections from the cortical larynx area in the squirrel monkey.</article-title> <source><italic>Exp. Brain Res.</italic></source> <volume>25</volume> <fpage>401</fpage>&#x2013;<lpage>411</lpage>. <pub-id pub-id-type="doi">10.1007/BF00241730</pub-id></citation></ref>
<ref id="B70"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>J&#x00FC;rgens</surname> <given-names>U.</given-names></name></person-group> (<year>1992</year>). <article-title>&#x201C;On the neurobiology of vocal communication,&#x201D; in</article-title> <source><italic>Nonverbal Vocal Communication: Comparative and Developmental Approaches</italic></source>, <role>eds</role> <person-group person-group-type="editor"><name><surname>Papousek</surname> <given-names>H.</given-names></name> <name><surname>J&#x00FC;rgens</surname> <given-names>U.</given-names></name> <name><surname>Papousek</surname> <given-names>M.</given-names></name></person-group> (<publisher-loc>Cambridge</publisher-loc>: <publisher-name>Cambridge University Press</publisher-name>), <fpage>31</fpage>&#x2013;<lpage>42</lpage>.</citation></ref>
<ref id="B71"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>J&#x00FC;rgens</surname> <given-names>U.</given-names></name></person-group> (<year>2002</year>). <article-title>Neural pathways underlying vocal control.</article-title> <source><italic>Neurosci. Biobehav. Rev.</italic></source> <volume>26</volume> <fpage>235</fpage>&#x2013;<lpage>258</lpage>. <pub-id pub-id-type="doi">10.1016/S0149-7634(01)00068-9</pub-id></citation></ref>
<ref id="B72"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Kegl</surname> <given-names>J.</given-names></name> <name><surname>Senghas</surname> <given-names>A.</given-names></name> <name><surname>Coppola</surname> <given-names>M.</given-names></name></person-group> (<year>1999</year>). <article-title>&#x201C;Creation through contact: sign language emergence and sign language change in nicaragua,&#x201D; in</article-title> <source><italic>Language Creation and Language Change: Creolization, Diachrony, and Development</italic></source>, <role>ed.</role> <person-group person-group-type="editor"><name><surname>DeGraff</surname> <given-names>M.</given-names></name></person-group> (<publisher-loc>Cambridge, MA</publisher-loc>: <publisher-name>MIT Press</publisher-name>), <fpage>179</fpage>&#x2013;<lpage>237</lpage>.</citation></ref>
<ref id="B73"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Kent</surname> <given-names>R. D.</given-names></name></person-group> (<year>1984</year>). <article-title>Psychobiology of speech development: coemergence of language and a movement system.</article-title> <source><italic>Am. J. Physiol.</italic></source> <volume>15</volume> <fpage>R888</fpage>&#x2013;<lpage>R894</lpage>. <pub-id pub-id-type="doi">10.1152/ajpregu.1984.246.6.R888</pub-id> <pub-id pub-id-type="pmid">6742163</pub-id></citation></ref>
<ref id="B74"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Kita</surname> <given-names>S.</given-names></name></person-group> (<year>1997</year>). <article-title>Two-dimensional semantic analysis of Japanese mimetics.</article-title> <source><italic>Linguistics</italic></source> <volume>35</volume> <fpage>379</fpage>&#x2013;<lpage>415</lpage>. <pub-id pub-id-type="doi">10.1515/ling.1997.35.2.379</pub-id></citation></ref>
<ref id="B75"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>K&#x00F6;hler</surname> <given-names>W.</given-names></name></person-group> (<year>1947</year>). <source><italic>Gestalt Psychology</italic></source>, <edition>2nd Edn</edition>. <publisher-loc>New York, NY</publisher-loc>: <publisher-name>Liveright Publishing</publisher-name>.</citation></ref>
<ref id="B76"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Kuypers</surname> <given-names>H. G.</given-names></name></person-group> (<year>1958</year>). <article-title>Corticobular connexions to the pons and lower brain-stem in man: an anatomical study.</article-title> <source><italic>Brain</italic></source> <volume>81</volume> <fpage>364</fpage>&#x2013;<lpage>388</lpage>. <pub-id pub-id-type="doi">10.1093/brain/81.3.364</pub-id> <pub-id pub-id-type="pmid">13596471</pub-id></citation></ref>
<ref id="B77"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Leavens</surname> <given-names>D.</given-names></name> <name><surname>Racine</surname> <given-names>T. P.</given-names></name> <name><surname>Hopkins</surname> <given-names>W. D.</given-names></name></person-group> (<year>2009</year>). <article-title>&#x201C;The ontogeny and phylogeny of non-verbal deixis,&#x201D; in</article-title> <source><italic>The Prehistory of Language</italic></source>, <role>eds</role> <person-group person-group-type="editor"><name><surname>Botha</surname> <given-names>R. P.</given-names></name> <name><surname>Knight</surname> <given-names>C.</given-names></name></person-group> (<publisher-loc>New York, NY</publisher-loc>: <publisher-name>Oxford University Press</publisher-name>), <fpage>142</fpage>&#x2013;<lpage>165</lpage>.</citation></ref>
<ref id="B78"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Lee</surname> <given-names>N.</given-names></name> <name><surname>Mikesell</surname> <given-names>L.</given-names></name> <name><surname>Joaquin</surname> <given-names>A. D. L.</given-names></name> <name><surname>Mates</surname> <given-names>A. W.</given-names></name> <name><surname>Schumann</surname> <given-names>J. H.</given-names></name></person-group> (<year>2009</year>). <source><italic>The Interactional Instinct: The Evolution and Acquisition of Language.</italic></source> <publisher-loc>New York, NY</publisher-loc>: <publisher-name>Oxford University Press</publisher-name>. <pub-id pub-id-type="doi">10.1093/acprof:oso/9780195384246.001.0001</pub-id></citation></ref>
<ref id="B79"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Lee</surname> <given-names>N.</given-names></name> <name><surname>Schumann</surname> <given-names>J. H.</given-names></name></person-group> (<year>2005</year>). <article-title>&#x201C;The interactional instinct: the evolution and acquisition of language,&#x201D; in</article-title> <source><italic>Paper Presented at the Congress of the International Association for Applied Linguistics</italic></source>, <publisher-loc>Madison, WI</publisher-loc>.</citation></ref>
<ref id="B80"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Lemon</surname> <given-names>R. N.</given-names></name></person-group> (<year>2008</year>). <article-title>Descending pathways in motor control.</article-title> <source><italic>Annu. Rev. Neurosci.</italic></source> <volume>31</volume> <fpage>195</fpage>&#x2013;<lpage>218</lpage>. <pub-id pub-id-type="doi">10.1146/annurev.neuro.31.060407.125547</pub-id></citation></ref>
<ref id="B81"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Lieberman</surname> <given-names>P.</given-names></name></person-group> (<year>1968</year>). <article-title>Primate vocalizations and human linguistic ability.</article-title> <source><italic>J. Acoust. Soc. Am.</italic></source> <volume>44</volume> <fpage>1574</fpage>&#x2013;<lpage>1584</lpage>. <pub-id pub-id-type="doi">10.1121/1.1911299</pub-id></citation></ref>
<ref id="B82"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Lieberman</surname> <given-names>P.</given-names></name></person-group> (<year>1984</year>). <source><italic>The Biology and Evolution of Language.</italic></source> <publisher-loc>Cambridge MA</publisher-loc>: <publisher-name>Harvard University Press</publisher-name>.</citation></ref>
<ref id="B83"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Lieberman</surname> <given-names>P.</given-names></name></person-group> (<year>2000</year>). <source><italic>Human Language and our Reptilian Brain.</italic></source> <publisher-loc>Cambridge MA</publisher-loc>: <publisher-name>Harvard University Press</publisher-name>.</citation></ref>
<ref id="B84"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Lieberman</surname> <given-names>P.</given-names></name></person-group> (<year>2003</year>). <article-title>&#x201C;Language evolution and innateness,&#x201D; in</article-title> <source><italic>Mind, Brain, and Language Multidisciplinary Perspectives</italic></source>, <role>eds</role> <person-group person-group-type="editor"><name><surname>Banich</surname> <given-names>M. T.</given-names></name> <name><surname>Mack</surname> <given-names>M.</given-names></name></person-group> (<publisher-loc>Mahwah, NJ</publisher-loc>: <publisher-name>Lawrence Erlbaum</publisher-name>), <fpage>3</fpage>&#x2013;<lpage>21</lpage>.</citation></ref>
<ref id="B85"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Lieberman</surname> <given-names>P.</given-names></name></person-group> (<year>2006a</year>). <article-title>Limits on tongue deformation: Diana monkey vocalizations and the impossible vocal tract shapes proposed by Riede et al. (2005).</article-title> <source><italic>J. Hum. Evol.</italic></source> <volume>50</volume> <fpage>219</fpage>&#x2013;<lpage>221</lpage>. <pub-id pub-id-type="doi">10.1016/j.jhevol.2005.07.010</pub-id> <pub-id pub-id-type="pmid">16376410</pub-id></citation></ref>
<ref id="B86"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Lieberman</surname> <given-names>P.</given-names></name></person-group> (<year>2006b</year>). <source><italic>Towards an Evolutionary Biology of Language.</italic></source> <publisher-loc>Cambridge, MA</publisher-loc>: <publisher-name>Harvard University Press</publisher-name>.</citation></ref>
<ref id="B87"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Lieberman</surname> <given-names>P.</given-names></name></person-group> (<year>2007</year>). <article-title>The evolution of human speech. Its anatomical and neural bases.</article-title> <source><italic>Curr. Anthropol.</italic></source> <volume>48</volume> <fpage>39</fpage>&#x2013;<lpage>66</lpage>. <pub-id pub-id-type="doi">10.1086/509092</pub-id> <pub-id pub-id-type="pmid">22106428</pub-id></citation></ref>
<ref id="B88"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Lieberman</surname> <given-names>P.</given-names></name></person-group> (<year>2016</year>). <article-title>The evolution of language and thought.</article-title> <source><italic>J. Anthropol. Sci.</italic></source> <volume>94</volume> <fpage>127</fpage>&#x2013;<lpage>146</lpage>. <pub-id pub-id-type="doi">10.4436/JASS.94029</pub-id> <pub-id pub-id-type="pmid">26963222</pub-id></citation></ref>
<ref id="B89"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Lieberman</surname> <given-names>P.</given-names></name> <name><surname>Crelin</surname> <given-names>E. S.</given-names></name> <name><surname>Klatt</surname> <given-names>D. H.</given-names></name></person-group> (<year>1972</year>). <article-title>Phonetic ability and related anatomy of the newborn and adult human, Neanderthal man, and the chimpanzee.</article-title> <source><italic>Am. Anthropol.</italic></source> <volume>74</volume> <fpage>287</fpage>&#x2013;<lpage>307</lpage>. <pub-id pub-id-type="doi">10.1525/aa.1972.74.3.02a00020</pub-id></citation></ref>
<ref id="B90"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Lieberman</surname> <given-names>P.</given-names></name> <name><surname>Laitman</surname> <given-names>J. T.</given-names></name> <name><surname>Reidenberg</surname> <given-names>J. S.</given-names></name> <name><surname>Gannon</surname> <given-names>P. J.</given-names></name></person-group> (<year>1992</year>). <article-title>The anatomy, physiology, acoustics and perception of speech: essential elements in analysis of the evolution of human speech.</article-title> <source><italic>J. Hum. Evol.</italic></source> <volume>23</volume> <fpage>447</fpage>&#x2013;<lpage>467</lpage>. <pub-id pub-id-type="doi">10.1016/0047-2484(92)90046-C</pub-id></citation></ref>
<ref id="B91"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Livingstone</surname> <given-names>M.</given-names></name> <name><surname>Srihasam</surname> <given-names>K.</given-names></name> <name><surname>Morocz</surname> <given-names>I.</given-names></name></person-group> (<year>2010</year>). <article-title>The benefit of symbols: monkeys show linear, human-like, accuracy when using symbols to represent scalar value.</article-title> <source><italic>Anim. Cogn.</italic></source> <volume>13</volume> <fpage>711</fpage>&#x2013;<lpage>719</lpage>. <pub-id pub-id-type="doi">10.1007/s10071-010-0321-1</pub-id> <pub-id pub-id-type="pmid">20443126</pub-id></citation></ref>
<ref id="B92"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Livingstone</surname> <given-names>M. S.</given-names></name> <name><surname>Pettine</surname> <given-names>W. W.</given-names></name> <name><surname>Srihasam</surname> <given-names>K.</given-names></name> <name><surname>Moore</surname> <given-names>B.</given-names></name> <name><surname>Morocz</surname> <given-names>I. A.</given-names></name> <name><surname>Lee</surname> <given-names>D.</given-names></name></person-group> (<year>2014</year>). <article-title>Symbol addition by monkeys provides evidence for normalized quantity coding.</article-title> <source><italic>Proc. Natl. Acad. Sci. U.S.A.</italic></source> <volume>111</volume> <fpage>6822</fpage>&#x2013;<lpage>6827</lpage>. <pub-id pub-id-type="doi">10.1073/pnas.1404208111</pub-id> <pub-id pub-id-type="pmid">24753600</pub-id></citation></ref>
<ref id="B93"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Locke</surname> <given-names>J. L.</given-names></name></person-group> (<year>1995</year>). <source><italic>The Child&#x2019;s Path to Spoken Language.</italic></source> <publisher-loc>Cambridge MA</publisher-loc>: <publisher-name>Harvard University Press</publisher-name>.</citation></ref>
<ref id="B94"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Lyn</surname> <given-names>H.</given-names></name> <name><surname>Savage-Rumbaugh</surname> <given-names>E. S.</given-names></name></person-group> (<year>2000</year>). <article-title>Observational word learning by two bonobos: ostensive and non-ostensive contexts.</article-title> <source><italic>Lang. Commun.</italic></source> <volume>20</volume> <fpage>255</fpage>&#x2013;<lpage>273</lpage>. <pub-id pub-id-type="doi">10.1016/S0271-5309(99)00026-9</pub-id></citation></ref>
<ref id="B95"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Lynch</surname> <given-names>M. P.</given-names></name> <name><surname>Oller</surname> <given-names>D. K.</given-names></name> <name><surname>Steffens</surname> <given-names>M. L.</given-names></name> <name><surname>Buder</surname> <given-names>E. H.</given-names></name></person-group> (<year>1995</year>). <article-title>Phrasing in prelinguistic vocalizations.</article-title> <source><italic>Dev. Psychobiol.</italic></source> <volume>28</volume> <fpage>3</fpage>&#x2013;<lpage>25</lpage>. <pub-id pub-id-type="doi">10.1002/dev.420280103</pub-id> <pub-id pub-id-type="pmid">7895922</pub-id></citation></ref>
<ref id="B96"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>MacKenzie</surname> <given-names>H.</given-names></name> <name><surname>Graham</surname> <given-names>S. A.</given-names></name> <name><surname>Curtin</surname> <given-names>S.</given-names></name></person-group> (<year>2011</year>). <article-title>Twelve-month-olds privilege words over other linguistic sounds in an associative learning task.</article-title> <source><italic>Dev. Sci.</italic></source> <volume>14</volume> <fpage>249</fpage>&#x2013;<lpage>255</lpage>. <pub-id pub-id-type="doi">10.1111/j.1467-7687.2010.00975.x</pub-id> <pub-id pub-id-type="pmid">22213898</pub-id></citation></ref>
<ref id="B97"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Maclarnon</surname> <given-names>A.</given-names></name> <name><surname>Hewitt</surname> <given-names>G.</given-names></name></person-group> (<year>1999</year>). <article-title>The evolution of human speech: the role of enhanced breathing control.</article-title> <source><italic>Am. J. Phys. Anthropol.</italic></source> <volume>109</volume> <fpage>341</fpage>&#x2013;<lpage>363</lpage>. <pub-id pub-id-type="doi">10.1002/(SICI)1096-8644(199907)109:3&#x003C;341::AID-AJPA5&#x003E;3.0.CO;2-2</pub-id> <pub-id pub-id-type="pmid">10407464</pub-id></citation></ref>
<ref id="B98"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Maclarnon</surname> <given-names>A.</given-names></name> <name><surname>Hewitt</surname> <given-names>G.</given-names></name></person-group> (<year>2004</year>). <article-title>Increased breathing control: another factor in the evolution of human language.</article-title> <source><italic>Evol. Anthropol. Issues News Rev.</italic></source> <volume>13</volume> <fpage>181</fpage>&#x2013;<lpage>197</lpage>. <pub-id pub-id-type="doi">10.1002/evan.20032</pub-id></citation></ref>
<ref id="B99"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>MacNeilage</surname> <given-names>P. F.</given-names></name></person-group> (<year>1998</year>). <article-title>The frame/content theory of evolution of speech production.</article-title> <source><italic>Behav. Brain Sci.</italic></source> <volume>21</volume> <fpage>499</fpage>&#x2013;<lpage>546</lpage>. <pub-id pub-id-type="doi">10.1017/S0140525X98001265</pub-id></citation></ref>
<ref id="B100"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>MacNeilage</surname> <given-names>P. F.</given-names></name></person-group> (<year>2008/2010</year>). <source><italic>The Origin of Speech.</italic></source> <publisher-loc>Oxford</publisher-loc>: <publisher-name>Oxford University Press</publisher-name>.</citation></ref>
<ref id="B101"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>MacNeilage</surname> <given-names>P. F.</given-names></name> <name><surname>Davis</surname> <given-names>B. L.</given-names></name></person-group> (<year>2000</year>). <article-title>On the origin of internal structure of word forms.</article-title> <source><italic>Science</italic></source> <volume>288</volume> <fpage>527</fpage>&#x2013;<lpage>531</lpage>. <pub-id pub-id-type="doi">10.1126/science.288.5465.527</pub-id> <pub-id pub-id-type="pmid">10775113</pub-id></citation></ref>
<ref id="B102"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>MacNeilage</surname> <given-names>P. F.</given-names></name> <name><surname>Davis</surname> <given-names>B. L.</given-names></name></person-group> (<year>2015</year>). <article-title>&#x201C;The evolution of language,&#x201D; in</article-title> <source><italic>The Handbook of Evolutionary Psychology</italic></source>, <role>ed.</role> <person-group person-group-type="editor"><name><surname>Buss</surname> <given-names>D. M.</given-names></name></person-group> (<publisher-loc>Hoboken, NJ</publisher-loc>: <publisher-name>Wiley</publisher-name>), <fpage>698</fpage>&#x2013;<lpage>723</lpage>.</citation></ref>
<ref id="B103"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Magrassi</surname> <given-names>L.</given-names></name> <name><surname>Aromataris</surname> <given-names>G.</given-names></name> <name><surname>Cabrini</surname> <given-names>A.</given-names></name> <name><surname>Annovazzi-Lodi</surname> <given-names>V.</given-names></name> <name><surname>Moro</surname> <given-names>A.</given-names></name></person-group> (<year>2015</year>). <article-title>Sound representation in higher language areas during language generation.</article-title> <source><italic>Proc. Natl. Acad. Sci. U.S.A.</italic></source> <volume>112</volume> <fpage>1868</fpage>&#x2013;<lpage>1873</lpage>. <pub-id pub-id-type="doi">10.1073/pnas.1418162112</pub-id> <pub-id pub-id-type="pmid">25624479</pub-id></citation></ref>
<ref id="B104"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Marno</surname> <given-names>H.</given-names></name> <name><surname>Farroni</surname> <given-names>T.</given-names></name> <name><surname>Dos Santos</surname> <given-names>Y. V.</given-names></name> <name><surname>Ekramnia</surname> <given-names>M.</given-names></name> <name><surname>Nespor</surname> <given-names>M.</given-names></name> <name><surname>Mehler</surname> <given-names>J.</given-names></name></person-group> (<year>2015</year>). <article-title>Can you see what I am talking about? Human speech triggers referential expectation in four-month-old infants.</article-title> <source><italic>Sci. Rep.</italic></source> <volume>5</volume>:<issue>13594</issue>. <pub-id pub-id-type="doi">10.1038/srep13594</pub-id> <pub-id pub-id-type="pmid">26323990</pub-id></citation></ref>
<ref id="B105"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Marno</surname> <given-names>H.</given-names></name> <name><surname>Guellai</surname> <given-names>B.</given-names></name> <name><surname>Vidal</surname> <given-names>Y.</given-names></name> <name><surname>Franzoi</surname> <given-names>J.</given-names></name> <name><surname>Nespor</surname> <given-names>M.</given-names></name> <name><surname>Mehler</surname> <given-names>J.</given-names></name></person-group> (<year>2016</year>). <article-title>Infants&#x2019; selectively pay attention to the information they receive from a native speaker of their language.</article-title> <source><italic>Front. Psychol.</italic></source> <volume>7</volume>:<issue>1150</issue>. <pub-id pub-id-type="doi">10.3389/fpsyg.2016.01150</pub-id> <pub-id pub-id-type="pmid">27536263</pub-id></citation></ref>
<ref id="B106"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Massaro</surname> <given-names>D. W.</given-names></name> <name><surname>Perlman</surname> <given-names>M.</given-names></name></person-group> (<year>2017</year>). <article-title>Quantifying iconicity&#x2019;s contribution during language acquisition: implications for vocabulary learning.</article-title> <source><italic>Front. Commun.</italic></source> <volume>2</volume>:<issue>4</issue>. <pub-id pub-id-type="doi">10.3389/fcomm.2017.00004</pub-id></citation></ref>
<ref id="B107"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Maurer</surname> <given-names>D.</given-names></name> <name><surname>Pathman</surname> <given-names>T.</given-names></name> <name><surname>Mondloch</surname> <given-names>C. J.</given-names></name></person-group> (<year>2006</year>). <article-title>The shape of boubas: sound-shape correspondences in toddlers and adults.</article-title> <source><italic>Dev. Sci.</italic></source> <volume>9</volume> <fpage>316</fpage>&#x2013;<lpage>322</lpage>. <pub-id pub-id-type="doi">10.1111/j.1467-7687.2006.00495.x</pub-id> <pub-id pub-id-type="pmid">16669803</pub-id></citation></ref>
<ref id="B108"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>McNeill</surname> <given-names>D.</given-names></name></person-group> (<year>2005</year>). <source><italic>Gesture and Thought.</italic></source> <publisher-loc>Chicago IL</publisher-loc>: <publisher-name>University of Chicago Press</publisher-name>. <pub-id pub-id-type="doi">10.7208/chicago/9780226514642.001.0001</pub-id></citation></ref>
<ref id="B109"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Monaghan</surname> <given-names>P.</given-names></name> <name><surname>Christiansen</surname> <given-names>M. H.</given-names></name></person-group> (<year>2006</year>). <article-title>&#x201C;Why form-meaning mappings are not entirely arbitrary in language,&#x201D; in</article-title> <source><italic>Proceedings of the 28th Annual Conference of the Cognitive Science Society</italic></source>, <role>ed.</role> <person-group person-group-type="editor"><name><surname>Sun</surname> <given-names>R.</given-names></name></person-group> (<publisher-loc>Hillsdale, NJ</publisher-loc>: <publisher-name>Lawrence Erlbaum</publisher-name>), <fpage>1838</fpage>&#x2013;<lpage>1843</lpage>.</citation></ref>
<ref id="B110"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Monaghan</surname> <given-names>P.</given-names></name> <name><surname>Christiansen</surname> <given-names>M. H.</given-names></name> <name><surname>Fitneva</surname> <given-names>S. A.</given-names></name></person-group> (<year>2011</year>). <article-title>The arbitrariness of the sign: learning advantages from the structure of the vocabulary.</article-title> <source><italic>J. Exp. Psychol. Gen.</italic></source> <volume>140</volume> <fpage>325</fpage>&#x2013;<lpage>347</lpage>. <pub-id pub-id-type="doi">10.1037/a0022924</pub-id> <pub-id pub-id-type="pmid">21517205</pub-id></citation></ref>
<ref id="B111"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Monaghan</surname> <given-names>P.</given-names></name> <name><surname>Mattock</surname> <given-names>K.</given-names></name> <name><surname>Walker</surname> <given-names>P.</given-names></name></person-group> (<year>2012</year>). <article-title>The role of sound symbolism in language learning.</article-title> <source><italic>J. Exp. Psychol. Mem. Cogn.</italic></source> <volume>38</volume> <fpage>1152</fpage>&#x2013;<lpage>1164</lpage>. <pub-id pub-id-type="doi">10.1037/a0027747</pub-id> <pub-id pub-id-type="pmid">22468804</pub-id></citation></ref>
<ref id="B112"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Monaghan</surname> <given-names>P.</given-names></name> <name><surname>Shillcock</surname> <given-names>R. C.</given-names></name> <name><surname>Christiansen</surname> <given-names>M. H.</given-names></name> <name><surname>Kirby</surname> <given-names>S.</given-names></name></person-group> (<year>2014</year>). <article-title>How arbitrary is language?</article-title> <source><italic>Philos. Trans. R. Soc. B Biol. Sci.</italic></source> <volume>369</volume>:<issue>20130299</issue>. <pub-id pub-id-type="doi">10.1098/rstb.2013.0299</pub-id> <pub-id pub-id-type="pmid">25092667</pub-id></citation></ref>
<ref id="B113"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Morton</surname> <given-names>E. S.</given-names></name></person-group> (<year>1994</year>). <article-title>&#x201C;Sound symbolism and its role in non-human vertebrate communication,&#x201D; in</article-title> <source><italic>Sound Symbolism</italic></source>, <role>eds</role> <person-group person-group-type="editor"><name><surname>Hinton</surname> <given-names>L.</given-names></name> <name><surname>Nichols</surname> <given-names>J.</given-names></name> <name><surname>Ohala</surname> <given-names>J. J.</given-names></name></person-group> (<publisher-loc>Cambridge</publisher-loc>: <publisher-name>Cambridge University Press</publisher-name>), <fpage>348</fpage>&#x2013;<lpage>365</lpage>.</citation></ref>
<ref id="B114"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Negus</surname> <given-names>V. E.</given-names></name></person-group> (<year>1949</year>). <source><italic>The Comparative Anatomy and Physiology of the Larynx.</italic></source> <publisher-loc>New York, NY</publisher-loc>: <publisher-name>Grune and Stratton</publisher-name>.</citation></ref>
<ref id="B115"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Nishimura</surname> <given-names>T.</given-names></name> <name><surname>Mikami</surname> <given-names>A.</given-names></name> <name><surname>Suzuki</surname> <given-names>J.</given-names></name> <name><surname>Matsuzawa</surname> <given-names>T.</given-names></name></person-group> (<year>2006</year>). <article-title>Descent of the hyoid in chimpanzees: evolution of face flattening and speech.</article-title> <source><italic>J. Hum. Evol.</italic></source> <volume>51</volume> <fpage>244</fpage>&#x2013;<lpage>254</lpage>. <pub-id pub-id-type="doi">10.1016/j.jhevol.2006.03.005</pub-id> <pub-id pub-id-type="pmid">16730049</pub-id></citation></ref>
<ref id="B116"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>N&#x00FA;&#x00F1;ez</surname> <given-names>R. E.</given-names></name></person-group> (<year>2017</year>). <article-title>Is there really an evolved capacity for number?</article-title> <source><italic>Trends Cogn. Sci.</italic></source> <volume>21</volume> <fpage>409</fpage>&#x2013;<lpage>424</lpage>. <pub-id pub-id-type="doi">10.1016/j.tics.2017.03.005</pub-id> <pub-id pub-id-type="pmid">28526128</pub-id></citation></ref>
<ref id="B117"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Ohala</surname> <given-names>J. J.</given-names></name></person-group> (<year>1994</year>). <article-title>&#x201C;The frequency code underlies the sound-symbolic use of voice pitch,&#x201D; in</article-title> <source><italic>Sound Symbolism</italic></source>, <role>eds</role> <person-group person-group-type="editor"><name><surname>Hinton</surname> <given-names>L.</given-names></name> <name><surname>Nichols</surname> <given-names>J.</given-names></name> <name><surname>Ohala</surname> <given-names>J. J.</given-names></name></person-group> (<publisher-loc>Cambridge</publisher-loc>: <publisher-name>Cambridge University Press</publisher-name>), <fpage>325</fpage>&#x2013;<lpage>365</lpage>.</citation></ref>
<ref id="B118"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Oller</surname> <given-names>D. K.</given-names></name></person-group> (<year>2000</year>). <source><italic>The Emergence of the Speech Capacity.</italic></source> <publisher-loc>Mahwah NJ</publisher-loc>: <publisher-name>Lawrence Erlbaum</publisher-name>. <pub-id pub-id-type="doi">10.4324/9781410602565</pub-id></citation></ref>
<ref id="B119"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Oller</surname> <given-names>D. K.</given-names></name> <name><surname>Griebel</surname> <given-names>U.</given-names></name></person-group> (<year>2014</year>). <article-title>On quantitative comparative research in communication and language evolution.</article-title> <source><italic>Biol. Theory</italic></source> <volume>9</volume> <fpage>296</fpage>&#x2013;<lpage>308</lpage>. <pub-id pub-id-type="doi">10.1007/s13752-014-0186-7</pub-id> <pub-id pub-id-type="pmid">25285057</pub-id></citation></ref>
<ref id="B120"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Oller</surname> <given-names>D. K.</given-names></name> <name><surname>Lynch</surname> <given-names>M. P.</given-names></name></person-group> (<year>1992</year>). <article-title>&#x201C;Infant vocalizations and innovations in infraphonology: toward a broader theory of development and disorders,&#x201D; in</article-title> <source><italic>Phonological Development: Models, Research, Implications</italic></source>, <role>eds</role> <person-group person-group-type="editor"><name><surname>Ferguson</surname> <given-names>C. A.</given-names></name> <name><surname>Menn</surname> <given-names>L.</given-names></name> <name><surname>Stoel-Gammon</surname> <given-names>C.</given-names></name></person-group> (<publisher-loc>Timonium, MD</publisher-loc>: <publisher-name>York Press</publisher-name>), <fpage>509</fpage>&#x2013;<lpage>536</lpage>.</citation></ref>
<ref id="B121"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Oudeyer</surname> <given-names>P.-Y.</given-names></name> <name><surname>Kaplan</surname> <given-names>F.</given-names></name></person-group> (<year>2006</year>). <article-title>Discovering communication.</article-title> <source><italic>Conn. Sci.</italic></source> <volume>18</volume> <fpage>189</fpage>&#x2013;<lpage>206</lpage>. <pub-id pub-id-type="doi">10.1080/09540090600768567</pub-id></citation></ref>
<ref id="B122"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Pedersen</surname> <given-names>J.</given-names></name></person-group> (<year>2012</year>). <article-title>&#x201C;The symbolic mind: apes, symbols, and the evolution of language,&#x201D; in</article-title> <source><italic>Paper Presented at the Graduate Theses and Dissertations</italic></source> <volume>12430</volume> (<publisher-loc>Ames, IA</publisher-loc>: <publisher-name>Iowa State University</publisher-name>).</citation></ref>
<ref id="B123"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Peirce</surname> <given-names>C. S.</given-names></name></person-group> (<year>1998</year>). <article-title>&#x201C;What is a sign?,&#x201D; in</article-title> <source><italic>The Essential Peirce: Selected Philosophical Writings</italic></source> <volume>Vol. 2</volume> <role>ed.</role> <person-group person-group-type="editor"><name><surname>Project</surname> <given-names>P. E.</given-names></name></person-group> (<publisher-loc>Bloomington IN</publisher-loc>: <publisher-name>Indiana University Press</publisher-name>), <fpage>1893</fpage>&#x2013;<lpage>1913</lpage>.</citation></ref>
<ref id="B124"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Penn</surname> <given-names>D. C.</given-names></name> <name><surname>Povinelli</surname> <given-names>D. J.</given-names></name></person-group> (<year>2007</year>). <article-title>On the lack of evidence that non-human animals possess anything remotely resembling a &#x2018;theory of mind&#x2019;.</article-title> <source><italic>Philos. Trans. R. Soc. B Biol. Sci.</italic></source> <volume>362</volume> <fpage>731</fpage>&#x2013;<lpage>744</lpage>. <pub-id pub-id-type="doi">10.1098/rstb.2006.2023</pub-id> <pub-id pub-id-type="pmid">17264056</pub-id></citation></ref>
<ref id="B125"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Perlman</surname> <given-names>M.</given-names></name> <name><surname>Dale</surname> <given-names>R.</given-names></name> <name><surname>Lupyan</surname> <given-names>G.</given-names></name></person-group> (<year>2015</year>). <article-title>Iconicity can ground the creation of vocal symbols.</article-title> <source><italic>R. Soc. Open Sci.</italic></source> <volume>2</volume>:<issue>150152</issue>. <pub-id pub-id-type="doi">10.1098/rsos.150152</pub-id> <pub-id pub-id-type="pmid">26361547</pub-id></citation></ref>
<ref id="B126"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Perruchet</surname> <given-names>P.</given-names></name> <name><surname>Peereman</surname> <given-names>R.</given-names></name> <name><surname>Tyler</surname> <given-names>M. D.</given-names></name></person-group> (<year>2006</year>). <article-title>Do we need algebraic-like computations? A reply to Bonatti, Pena, Nespor, and Mehler (2006).</article-title> <source><italic>J. Exp. Psychol. Gen.</italic></source> <volume>135</volume> <fpage>322</fpage>&#x2013;<lpage>326</lpage>. <pub-id pub-id-type="doi">10.1037/0096-3445.135.2.322</pub-id></citation></ref>
<ref id="B127"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Perruchet</surname> <given-names>P.</given-names></name> <name><surname>Rey</surname> <given-names>A.</given-names></name></person-group> (<year>2005</year>). <article-title>Does the mastery of center-embedded linguistic structures distinguish humans from nonhuman primates?</article-title> <source><italic>Psychon. Bull. Rev.</italic></source> <volume>12</volume> <fpage>307</fpage>&#x2013;<lpage>313</lpage>. <pub-id pub-id-type="doi">10.3758/BF03196377</pub-id> <pub-id pub-id-type="pmid">16082811</pub-id></citation></ref>
<ref id="B128"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Perruchet</surname> <given-names>P.</given-names></name> <name><surname>Tyler</surname> <given-names>M. D.</given-names></name> <name><surname>Galland</surname> <given-names>N.</given-names></name> <name><surname>Peereman</surname> <given-names>R.</given-names></name></person-group> (<year>2004</year>). <article-title>Learning nonadjacent dependencies: no need for algebraic-like computations.</article-title> <source><italic>J. Exp. Psychol. Gen.</italic></source> <volume>133</volume> <fpage>573</fpage>&#x2013;<lpage>583</lpage>. <pub-id pub-id-type="doi">10.1037/0096-3445.133.4.573</pub-id> <pub-id pub-id-type="pmid">15584807</pub-id></citation></ref>
<ref id="B129"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Pika</surname> <given-names>S.</given-names></name> <name><surname>Liebal</surname> <given-names>K.</given-names></name> <name><surname>Call</surname> <given-names>J.</given-names></name> <name><surname>Tomasello</surname> <given-names>M.</given-names></name></person-group> (<year>2005</year>). <article-title>The gestural communication of apes.</article-title> <source><italic>Gesture</italic></source> <volume>5</volume> <fpage>41</fpage>&#x2013;<lpage>56</lpage>. <pub-id pub-id-type="doi">10.1075/gest.5.1-2.05pik</pub-id></citation></ref>
<ref id="B130"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Pinker</surname> <given-names>S.</given-names></name></person-group> (<year>1994</year>). <source><italic>The Language Instinct.</italic></source> <publisher-loc>New York, NY</publisher-loc>: <publisher-name>William Morrow</publisher-name>.</citation></ref>
<ref id="B131"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Pinker</surname> <given-names>S.</given-names></name> <name><surname>Jackendoff</surname> <given-names>R.</given-names></name></person-group> (<year>2005</year>). <article-title>The faculty of language: what&#x2019;s special about it?</article-title> <source><italic>Cognition</italic></source> <volume>95</volume> <fpage>201</fpage>&#x2013;<lpage>236</lpage>. <pub-id pub-id-type="doi">10.1016/j.cognition.2004.08.004</pub-id> <pub-id pub-id-type="pmid">15694646</pub-id></citation></ref>
<ref id="B132"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Pleyer</surname> <given-names>M.</given-names></name></person-group> (<year>2017</year>). <article-title>Protolanguage and mechanisms of meaning construal in interaction.</article-title> <source><italic>Lang. Sci.</italic></source> <volume>63</volume> <fpage>69</fpage>&#x2013;<lpage>90</lpage>. <pub-id pub-id-type="doi">10.1016/j.langsci.2017.01.003</pub-id></citation></ref>
<ref id="B133"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Pleyer</surname> <given-names>M.</given-names></name> <name><surname>Winters</surname> <given-names>J.</given-names></name></person-group> (<year>2015</year>). <article-title>Integrating cognitive linguistics and language evolution research.</article-title> <source><italic>Theoria Hist. Sci.</italic></source> <volume>11</volume> <fpage>19</fpage>&#x2013;<lpage>43</lpage>. <pub-id pub-id-type="doi">10.12775/ths-2014-002</pub-id> <pub-id pub-id-type="pmid">26988387</pub-id></citation></ref>
<ref id="B134"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Pollick</surname> <given-names>A. S.</given-names></name> <name><surname>de Waal</surname> <given-names>F. B. M.</given-names></name></person-group> (<year>2007</year>). <article-title>Ape gestures and language evolution.</article-title> <source><italic>Proc. Natl. Acad. Sci. U.S.A.</italic></source> <volume>104</volume> <fpage>8184</fpage>&#x2013;<lpage>8189</lpage>. <pub-id pub-id-type="doi">10.1073/pnas.0702624104</pub-id> <pub-id pub-id-type="pmid">17470779</pub-id></citation></ref>
<ref id="B135"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Price</surname> <given-names>T.</given-names></name> <name><surname>Wadewitz</surname> <given-names>P.</given-names></name> <name><surname>Cheney</surname> <given-names>D.</given-names></name> <name><surname>Seyfarth</surname> <given-names>R.</given-names></name> <name><surname>Hammerschmidt</surname> <given-names>K.</given-names></name> <name><surname>Fischer</surname> <given-names>J.</given-names></name></person-group> (<year>2015</year>). <article-title>Vervets revisited: a quantitative analysis of alarm call structure and context specificity.</article-title> <source><italic>Sci. Rep.</italic></source> <volume>5</volume>:<issue>13220</issue>. <pub-id pub-id-type="doi">10.1038/srep13220</pub-id> <pub-id pub-id-type="pmid">26286236</pub-id></citation></ref>
<ref id="B136"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Provine</surname> <given-names>R. R.</given-names></name></person-group> (<year>2005</year>). <article-title>Walkie-talkie evolution: bipedalism and vocal production.</article-title> <source><italic>Behav. Brain Sci.</italic></source> <volume>27</volume> <fpage>520</fpage>&#x2013;<lpage>521</lpage>. <pub-id pub-id-type="doi">10.1017/S0140525X04410115</pub-id></citation></ref>
<ref id="B137"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Rizzolatti</surname> <given-names>G.</given-names></name> <name><surname>Arbib</surname> <given-names>M. A.</given-names></name></person-group> (<year>1998</year>). <article-title>Language within our grasp.</article-title> <source><italic>Trends Neurosci.</italic></source> <volume>21</volume> <fpage>188</fpage>&#x2013;<lpage>194</lpage>. <pub-id pub-id-type="doi">10.1016/S0166-2236(98)01260-0</pub-id></citation></ref>
<ref id="B138"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Sandler</surname> <given-names>W.</given-names></name></person-group> (<year>2012</year>). <source><italic>Dedicated Gestures and the Emergence of Sign Language.</italic></source> <fpage>265</fpage>&#x2013;<lpage>307</lpage>. Available at: <ext-link ext-link-type="uri" xlink:href="http://sandlersignlab.haifa.ac.il/html/html_eng/Dedicated_Gestures.pdf">http://sandlersignlab.haifa.ac.il/html/html_eng/Dedicated_Gestures.pdf</ext-link></citation></ref>
<ref id="B139"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Sandler</surname> <given-names>W.</given-names></name></person-group> (<year>2013</year>). <article-title>Vive la diff&#x00E9;rence: sign language and spoken language in language evolution.</article-title> <source><italic>Lang. Cogn.</italic></source> <volume>5</volume> <fpage>189</fpage>&#x2013;<lpage>203</lpage>. <pub-id pub-id-type="doi">10.1515/langcog-2013-0013</pub-id></citation></ref>
<ref id="B140"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Sandler</surname> <given-names>W.</given-names></name></person-group> (<year>2017</year>). <article-title>The challenge of sign language phonology.</article-title> <source><italic>Annu. Rev. Linguist.</italic></source> <volume>7</volume> <fpage>43</fpage>&#x2013;<lpage>63</lpage>. <pub-id pub-id-type="doi">10.3758/s13428-015-0579-y</pub-id> <pub-id pub-id-type="pmid">25939978</pub-id></citation></ref>
<ref id="B141"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Sandler</surname> <given-names>W.</given-names></name> <name><surname>Aronoff</surname> <given-names>M.</given-names></name> <name><surname>Padden</surname> <given-names>C.</given-names></name> <name><surname>Meir</surname> <given-names>I.</given-names></name></person-group> (<year>2014</year>). &#x201C;<article-title>Language emergence Al-sayyid bedouin sign language</article-title>,&#x201D; in <source><italic>The Cambridge Handbook of Linguistic Anthropology</italic></source>, <role>eds</role> <person-group person-group-type="editor"><name><surname>Enfield</surname> <given-names>N. J.</given-names></name> <name><surname>Kockelman</surname> <given-names>P.</given-names></name> <name><surname>Sidnell</surname> <given-names>J.</given-names></name></person-group> (<publisher-loc>Cambridge</publisher-loc>: <publisher-name>Cambridge University Press</publisher-name>), <fpage>246</fpage>&#x2013;<lpage>278</lpage>.</citation></ref>
<ref id="B142"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Sapir</surname> <given-names>E.</given-names></name></person-group> (<year>1929</year>). <article-title>A study in phonetic symbolism.</article-title> <source><italic>J. Exp. Psychol.</italic></source> <volume>12</volume> <fpage>225</fpage>&#x2013;<lpage>239</lpage>. <pub-id pub-id-type="doi">10.1037/h0070931</pub-id></citation></ref>
<ref id="B143"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Sasaki</surname> <given-names>S.</given-names></name> <name><surname>Isa</surname> <given-names>T.</given-names></name> <name><surname>Pettersson</surname> <given-names>L.-G.</given-names></name> <name><surname>Alstermark</surname> <given-names>B.</given-names></name> <name><surname>Naito</surname> <given-names>K.</given-names></name> <name><surname>Yoshimura</surname> <given-names>K.</given-names></name><etal/></person-group> (<year>2004</year>). <article-title>Dexterous finger movements in primate without monosynaptic corticomotoneuronal excitation.</article-title> <source><italic>J. Neurophysiol.</italic></source> <volume>92</volume> <fpage>3142</fpage>&#x2013;<lpage>3147</lpage>. <pub-id pub-id-type="doi">10.1152/jn.00342.2004</pub-id> <pub-id pub-id-type="pmid">15175371</pub-id></citation></ref>
<ref id="B144"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Saussure</surname> <given-names>F. D.</given-names></name></person-group> (<year>1916/1966</year>). <source><italic>Course in General Linguistics.</italic></source> <publisher-loc>New York, NY</publisher-loc>: <publisher-name>McGraw-Hill</publisher-name>.</citation></ref>
<ref id="B145"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Savage-Rumbaugh</surname> <given-names>E. S.</given-names></name> <name><surname>Fields</surname> <given-names>W.</given-names></name></person-group> (<year>2000</year>). <article-title>Linguistic, cultural and cognitive capacities of bonobos (<italic>Pan paniscus</italic>).</article-title> <source><italic>Cult. Psychol.</italic></source> <volume>6</volume> <fpage>131</fpage>&#x2013;<lpage>153</lpage>. <pub-id pub-id-type="doi">10.1177/1354067X0062003</pub-id></citation></ref>
<ref id="B146"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Savage-Rumbaugh</surname> <given-names>E. S.</given-names></name> <name><surname>McDonald</surname> <given-names>K.</given-names></name> <name><surname>Sevcik</surname> <given-names>R. A.</given-names></name> <name><surname>Hopkins</surname> <given-names>W. D.</given-names></name> <name><surname>Rubert</surname> <given-names>E.</given-names></name></person-group> (<year>1986</year>). <article-title>Spontaneous symbol acquisition and communicative use by pygmy chimpanzees (<italic>Pan paniscus</italic>).</article-title> <source><italic>J. Exp. Psychol. Gen.</italic></source> <volume>115</volume> <fpage>211</fpage>&#x2013;<lpage>235</lpage>. <pub-id pub-id-type="doi">10.1037/0096-3445.115.3.211</pub-id> <pub-id pub-id-type="pmid">2428917</pub-id></citation></ref>
<ref id="B147"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Savage-Rumbaugh</surname> <given-names>E. S.</given-names></name> <name><surname>Murphy</surname> <given-names>J.</given-names></name> <name><surname>Sevcik</surname> <given-names>R. A.</given-names></name> <name><surname>Brakke</surname> <given-names>K. E.</given-names></name> <name><surname>Williams</surname> <given-names>S. L.</given-names></name> <name><surname>Rumbaugh</surname> <given-names>D. M.</given-names></name></person-group> (<year>1993</year>). <article-title>Language comprehension in ape and child.</article-title> <source><italic>Monogr. Soc. Res. Child Dev.</italic></source> <volume>58</volume> <fpage>1</fpage>&#x2013;<lpage>221</lpage>. <pub-id pub-id-type="doi">10.2307/1166068</pub-id></citation></ref>
<ref id="B148"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Savage-Rumbaugh</surname> <given-names>E. S.</given-names></name> <name><surname>Rumbaugh</surname> <given-names>D. M.</given-names></name></person-group> (<year>1978</year>). <article-title>Symbolization, language, and chimpanzees: a theoretical reevaluation based on initial language acquisition processes in four young <italic>Pan troglodytes</italic>.</article-title> <source><italic>Brain Lang.</italic></source> <volume>6</volume> <fpage>265</fpage>&#x2013;<lpage>300</lpage>. <pub-id pub-id-type="doi">10.1016/0093-934X(78)90063-9</pub-id> <pub-id pub-id-type="pmid">737530</pub-id></citation></ref>
<ref id="B149"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Savage-Rumbaugh</surname> <given-names>S.</given-names></name> <name><surname>Shanker</surname> <given-names>S. G.</given-names></name> <name><surname>Taylor</surname> <given-names>T. J.</given-names></name></person-group> (<year>2001</year>). <source><italic>Apes, Language, and the Human Mind.</italic></source> <publisher-loc>New York, NY</publisher-loc>: <publisher-name>Oxford University Press</publisher-name>.</citation></ref>
<ref id="B150"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Savage-Rumbaugh</surname> <given-names>S. E.</given-names></name> <name><surname>Lewin</surname> <given-names>R.</given-names></name></person-group> (<year>1996</year>). <source><italic>Kanzi: The ape at the Brink of the Human Mind.</italic></source> <publisher-loc>New York, NY</publisher-loc>: <publisher-name>Wiley</publisher-name>.</citation></ref>
<ref id="B151"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Senghas</surname> <given-names>A.</given-names></name></person-group> (<year>1995</year>). <source><italic>Children&#x2019;s Contribution to the Birth of Nicaraguan Sign Language.</italic></source> <comment>Doctoral dissertation</comment>, <publisher-loc>Massachusetts Institute of Technology, Boston, MA</publisher-loc>.</citation></ref>
<ref id="B152"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Sereno</surname> <given-names>M. I.</given-names></name></person-group> (<year>2014</year>). <article-title>Origin of symbol-using systems: speech, but not sign, without the semantic urge.</article-title> <source><italic>Philoso. Trans. R. Soc. B Biol. Sci.</italic></source> <volume>369</volume>:<issue>20130303</issue>. <pub-id pub-id-type="doi">10.1098/rstb.2013.0303</pub-id> <pub-id pub-id-type="pmid">25092671</pub-id></citation></ref>
<ref id="B153"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Seyfarth</surname> <given-names>R. M.</given-names></name> <name><surname>Cheney</surname> <given-names>D. L.</given-names></name> <name><surname>Marler</surname> <given-names>P.</given-names></name></person-group> (<year>1980</year>). <article-title>Vervet monkey alarm calls: semantic communication in a free-ranging primate.</article-title> <source><italic>Anim. Behav.</italic></source> <volume>28</volume> <fpage>1070</fpage>&#x2013;<lpage>1094</lpage>. <pub-id pub-id-type="doi">10.1016/S0003-3472(80)80097-2</pub-id></citation></ref>
<ref id="B154"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Simonyan</surname> <given-names>K.</given-names></name></person-group> (<year>2014</year>). <article-title>The laryngeal motor cortex: its organization and connectivity.</article-title> <source><italic>Curr. Opin. Neurobiol.</italic></source> <volume>28</volume> <fpage>15</fpage>&#x2013;<lpage>21</lpage>. <pub-id pub-id-type="doi">10.1016/j.conb.2014.05.006</pub-id> <pub-id pub-id-type="pmid">24929930</pub-id></citation></ref>
<ref id="B155"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Simonyan</surname> <given-names>K.</given-names></name> <name><surname>Horwitz</surname> <given-names>B.</given-names></name></person-group> (<year>2011</year>). <article-title>Laryngeal motor cortex and control of speech in humans.</article-title> <source><italic>Neuroscientist</italic></source> <volume>17</volume> <fpage>197</fpage>&#x2013;<lpage>208</lpage>. <pub-id pub-id-type="doi">10.1177/1073858410386727</pub-id> <pub-id pub-id-type="pmid">21362688</pub-id></citation></ref>
<ref id="B156"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Snodgrass</surname> <given-names>J. G.</given-names></name> <name><surname>Vanderwart</surname> <given-names>M.</given-names></name></person-group> (<year>1980</year>). <article-title>A standardized set of 260 pictures: norms for name agreement, image agreement, familiarity, and visual complexity.</article-title> <source><italic>J. Exp. Psychol. Hum. Learn. Mem.</italic></source> <volume>6</volume> <fpage>174</fpage>&#x2013;<lpage>215</lpage>. <pub-id pub-id-type="doi">10.1037/0278-7393.6.2.174</pub-id></citation></ref>
<ref id="B157"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Sonnweber</surname> <given-names>R.</given-names></name> <name><surname>Ravignani</surname> <given-names>A.</given-names></name> <name><surname>Fitch</surname> <given-names>W. T.</given-names></name></person-group> (<year>2015</year>). <article-title>Non-adjacent visual dependency learning in chimpanzees.</article-title> <source><italic>Anim. Cogn.</italic></source> <volume>18</volume> <fpage>733</fpage>&#x2013;<lpage>745</lpage>. <pub-id pub-id-type="doi">10.1007/s10071-015-0840-x</pub-id> <pub-id pub-id-type="pmid">25604423</pub-id></citation></ref>
<ref id="B158"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Squire</surname> <given-names>L. R.</given-names></name> <name><surname>Zola</surname> <given-names>S. M.</given-names></name></person-group> (<year>1996</year>). <article-title>Structure and function of declarative and nondeclarative memory systems.</article-title> <source><italic>Proc. Natl. Acad. Sci. U.S.A.</italic></source> <volume>93</volume> <fpage>13515</fpage>&#x2013;<lpage>13522</lpage>. <pub-id pub-id-type="doi">10.1073/pnas.93.24.13515</pub-id></citation></ref>
<ref id="B159"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Stjernfelt</surname> <given-names>F.</given-names></name></person-group> (<year>2012</year>). &#x201C;<article-title>The evolution of semiotic self-control</article-title>,&#x201D; in <source><italic>The Symbolic Species Evolved</italic></source>, <role>eds</role> <person-group person-group-type="editor"><name><surname>Schilhab</surname> <given-names>T.</given-names></name> <name><surname>Stjernfelt</surname> <given-names>F.</given-names></name> <name><surname>Deacon</surname> <given-names>T. W.</given-names></name></person-group> (<publisher-loc>Dordrecht</publisher-loc>: <publisher-name>Springer</publisher-name>), <fpage>39</fpage>&#x2013;<lpage>63</lpage>. <pub-id pub-id-type="doi">10.1007/978-94-007-2336-8_3</pub-id> <pub-id pub-id-type="pmid">10818558</pub-id></citation></ref>
<ref id="B160"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Studdert-Kennedy</surname> <given-names>M.</given-names></name></person-group> (<year>1998</year>). &#x201C;<article-title>Introduction: the emergence of phonology</article-title>,&#x201D; in <source><italic>Approaches to the Evolution of Language: Social and Cognitive Bases</italic></source>, <role>eds</role> <person-group person-group-type="editor"><name><surname>Hurford</surname> <given-names>J. R.</given-names></name> <name><surname>Studdert-Kennedy</surname> <given-names>M.</given-names></name> <name><surname>Knight</surname> <given-names>C.</given-names></name></person-group> (<publisher-loc>Cambridge</publisher-loc>: <publisher-name>Cambridge University Press</publisher-name>).</citation></ref>
<ref id="B161"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Sutton</surname> <given-names>D.</given-names></name> <name><surname>Larson</surname> <given-names>C.</given-names></name> <name><surname>Taylor</surname> <given-names>E. M.</given-names></name> <name><surname>Lindeman</surname> <given-names>R. C.</given-names></name></person-group> (<year>1973</year>). <article-title>Vocalization in rhesus monkeys: conditionability.</article-title> <source><italic>Brain Res.</italic></source> <volume>52</volume> <fpage>225</fpage>&#x2013;<lpage>231</lpage>. <pub-id pub-id-type="doi">10.1016/0006-8993(73)90660-4</pub-id> <pub-id pub-id-type="pmid">4633714</pub-id></citation></ref>
<ref id="B162"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Thom</surname> <given-names>S. A.</given-names></name> <name><surname>Hoit</surname> <given-names>J. D.</given-names></name> <name><surname>Hixon</surname> <given-names>T. J.</given-names></name> <name><surname>Smith</surname> <given-names>A. E.</given-names></name></person-group> (<year>2006</year>). <article-title>Velopharyngeal function during vocalization in infants.</article-title> <source><italic>Cleft Palate Craniofac. J.</italic></source> <volume>43</volume> <fpage>539</fpage>&#x2013;<lpage>546</lpage>. <pub-id pub-id-type="doi">10.1597/05-113</pub-id> <pub-id pub-id-type="pmid">16986991</pub-id></citation></ref>
<ref id="B163"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Thompson</surname> <given-names>R. L.</given-names></name> <name><surname>Vinson</surname> <given-names>D. P.</given-names></name> <name><surname>Woll</surname> <given-names>B.</given-names></name> <name><surname>Vigliocco</surname> <given-names>G.</given-names></name></person-group> (<year>2012</year>). <article-title>The road to language learning is iconic: evidence from British Sign Language.</article-title> <source><italic>Psychol. Sci.</italic></source> <volume>23</volume> <fpage>1443</fpage>&#x2013;<lpage>1448</lpage>. <pub-id pub-id-type="doi">10.1177/0956797612459763</pub-id> <pub-id pub-id-type="pmid">23150275</pub-id></citation></ref>
<ref id="B164"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Tomasello</surname> <given-names>M.</given-names></name></person-group> (<year>1990</year>). &#x201C;<article-title>Cultural transmission in the tool use and communicatory signaling of chimpanzees?</article-title>,&#x201D; in <source><italic>Language&#x2019; and Intelligence in Monkeys and Apes: Comparative Developmental Perspectives</italic></source>, <role>eds</role> <person-group person-group-type="editor"><name><surname>Parker</surname> <given-names>S. T.</given-names></name> <name><surname>Gibson</surname> <given-names>K. R.</given-names></name></person-group> (<publisher-loc>New York, NY</publisher-loc>: <publisher-name>Cambridge University Press</publisher-name>), <fpage>274</fpage>&#x2013;<lpage>311</lpage>.</citation></ref>
<ref id="B165"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Tomasello</surname> <given-names>M.</given-names></name></person-group> (<year>1996</year>). &#x201C;<article-title>Do apes ape?</article-title>,&#x201D; in <source><italic>Social Learning in Animals: the Roots of Culture</italic></source>, <role>eds</role> <person-group person-group-type="editor"><name><surname>Heyes</surname> <given-names>C. M.</given-names></name> <name><surname>Galef</surname> <given-names>B. G.</given-names></name></person-group> (<publisher-loc>San Diego, CA</publisher-loc>: <publisher-name>Academic Press</publisher-name>), <fpage>319</fpage>&#x2013;<lpage>346</lpage>. <pub-id pub-id-type="doi">10.1016/B978-012273965-1/50016-9</pub-id></citation></ref>
<ref id="B166"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Tomasello</surname> <given-names>M.</given-names></name> <name><surname>Zuberb&#x00FC;hler</surname> <given-names>K.</given-names></name></person-group> (<year>2002</year>). &#x201C;<article-title>Primate vocal and gestural communication</article-title>,&#x201D; in <source><italic>The Cognitive Animal: Empirical and Theoretical Perspectives on Animal Cognition</italic></source>, <role>eds</role> <person-group person-group-type="editor"><name><surname>Bekoff</surname> <given-names>M.</given-names></name> <name><surname>Allen</surname> <given-names>C.</given-names></name> <name><surname>Burghardt</surname> <given-names>G. M.</given-names></name></person-group> (<publisher-loc>Cambridge, MA</publisher-loc>: <publisher-name>MIT Press</publisher-name>), <fpage>293</fpage>&#x2013;<lpage>299</lpage>.</citation></ref>
<ref id="B167"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Vale</surname> <given-names>G. L.</given-names></name> <name><surname>Davis</surname> <given-names>S. J.</given-names></name> <name><surname>Lambeth</surname> <given-names>S. P.</given-names></name> <name><surname>Schapiro</surname> <given-names>S. J.</given-names></name> <name><surname>Whiten</surname> <given-names>A.</given-names></name></person-group> (<year>2017</year>). <article-title>Acquisition of a socially learned tool use sequence in chimpanzees: implications for cumulative culture.</article-title> <source><italic>Evol. Hum. Behav.</italic></source> <volume>38</volume> <fpage>635</fpage>&#x2013;<lpage>644</lpage>. <pub-id pub-id-type="doi">10.1016/j.evolhumbehav.2017.04.007</pub-id> <pub-id pub-id-type="pmid">29333058</pub-id></citation></ref>
<ref id="B168"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Vale</surname> <given-names>G. L.</given-names></name> <name><surname>Flynn</surname> <given-names>E. G.</given-names></name> <name><surname>Pender</surname> <given-names>L.</given-names></name> <name><surname>Price</surname> <given-names>E.</given-names></name> <name><surname>Whiten</surname> <given-names>A.</given-names></name> <name><surname>Lambeth</surname> <given-names>S. P.</given-names></name><etal/></person-group> (<year>2016</year>). <article-title>Robust retention and transfer of tool construction techniques in chimpanzees (<italic>Pan troglodytes</italic>).</article-title> <source><italic>J. Comp. Psychol.</italic></source> <volume>130</volume> <fpage>24</fpage>&#x2013;<lpage>35</lpage>. <pub-id pub-id-type="doi">10.1037/a0040000</pub-id> <pub-id pub-id-type="pmid">26881941</pub-id></citation></ref>
<ref id="B169"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Walker</surname> <given-names>P.</given-names></name> <name><surname>Bremner</surname> <given-names>J. G.</given-names></name> <name><surname>Mason</surname> <given-names>U.</given-names></name> <name><surname>Spring</surname> <given-names>J.</given-names></name> <name><surname>Mattock</surname> <given-names>K.</given-names></name> <name><surname>Slater</surname> <given-names>A.</given-names></name><etal/></person-group> (<year>2010</year>). <article-title>Preverbal infants&#x2019; sensitivity to synaesthetic cross-modality correspondences.</article-title> <source><italic>Psychol. Sci.</italic></source> <volume>21</volume> <fpage>21</fpage>&#x2013;<lpage>25</lpage>. <pub-id pub-id-type="doi">10.1177/0956797609354734</pub-id> <pub-id pub-id-type="pmid">20424017</pub-id></citation></ref>
<ref id="B170"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Wechsler</surname> <given-names>D.</given-names></name></person-group> (<year>1997</year>). <source><italic>WAIS-III, Wechsler adult Intelligence Scale.</italic></source> <publisher-loc>San Antonio TX</publisher-loc>: <publisher-name>Psychological Corporation</publisher-name>.</citation></ref>
<ref id="B171"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Whiten</surname> <given-names>A.</given-names></name> <name><surname>McGuigan</surname> <given-names>N.</given-names></name> <name><surname>Marshall-Pescini</surname> <given-names>S.</given-names></name> <name><surname>Hopper</surname> <given-names>L. M.</given-names></name></person-group> (<year>2009</year>). <article-title>Emulation, imitation, over-imitation and the scope of culture for child and chimpanzee.</article-title> <source><italic>Philos. Trans. R. Soc. B Biol. Sci.</italic></source> <volume>364</volume> <fpage>2417</fpage>&#x2013;<lpage>2428</lpage>. <pub-id pub-id-type="doi">10.1098/rstb.2009.0069</pub-id> <pub-id pub-id-type="pmid">19620112</pub-id></citation></ref>
<ref id="B172"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Williams</surname> <given-names>P.</given-names></name> <name><surname>Beer</surname> <given-names>R.</given-names></name> <name><surname>Gasser</surname> <given-names>M.</given-names></name></person-group> (<year>2008</year>). &#x201C;<article-title>Evolving referential communication in embodied dynamical agents</article-title>,&#x201D; in <source><italic>Paper Presented at the International Conference on the Simulation and Synthesis of Living Systems</italic></source>, (<publisher-loc>Cambridge, MA</publisher-loc>: <publisher-name>MIT Press</publisher-name>).</citation></ref>
<ref id="B173"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Wirth</surname> <given-names>S.</given-names></name> <name><surname>Yanike</surname> <given-names>M.</given-names></name> <name><surname>Frank</surname> <given-names>L. M.</given-names></name> <name><surname>Smith</surname> <given-names>A. C.</given-names></name> <name><surname>Brown</surname> <given-names>E. N.</given-names></name> <name><surname>Suzuki</surname> <given-names>W. A.</given-names></name></person-group> (<year>2003</year>). <article-title>Single neurons in the monkey hippocampus and learning of new associations.</article-title> <source><italic>Science</italic></source> <volume>300</volume> <fpage>1578</fpage>&#x2013;<lpage>1581</lpage>. <pub-id pub-id-type="doi">10.1126/science.1084324</pub-id> <pub-id pub-id-type="pmid">12791995</pub-id></citation></ref>
</ref-list>
</back>
</article>