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What can metacognition teach us about the evolution of communication?
Edited by Joëlle Proust
[Evolutionary Linguistic Theory 5:1] 2023
► pp. 11–52

Informational communication and metacognition

 | Institut Jean-NicodParis
Procedural metacognition is the set of affect-based mechanisms allowing agents to regulate cognitive actions like perceptual discrimination, memory retrieval or problem solving. This article proposes that procedural metacognition has had a major role in the evolution of communication. A plausible hypothesis is that, under pressure for maximizing signalling efficiency, the metacognitive abilities used by nonhumans to regulate their perception and their memory have been re-used to regulate their communication. On this view, detecting one’s production errors in signalling, or solving species-specific trade-offs between informativeness, processing effort, clarity, or urgency depend on a form of procedural metacognition, called “metacommunication”. How does this view relate to Gricean theories of human communication? A parallel between procedural trade-offs and conversational maxims is discussed for its evolutionary implications. Rather than accepting radically discontinuist interpretations, in which mindreading operates a full reorganization of pragmatics, it is proposed that procedural forms of regulation are entrenched in all forms of human communication. According to contextual demands, humans adopt and monitor more or less demanding informational goals, such as factual updating, clarifying, explaining, proving, and reaching consensus in collective matters. Under time pressure, only part of these goals require adopting others’ viewpoint. Efficiency in collective decision-making, in particular, might have been considerably raised by an ability to interpret others’ intentions and motivations.
Keywords: evolution of communication, animal signalling, procedural metacognition, cognitive goals, relevance, ostension, signalling trade-offs, informativeness, effortfulness, deception
Article outline
  • Introduction
    • From information to communication
    • How to define communication?
    • Metacommunication belongs to procedural metacognition
  • I.Metacognition in nonhumans
    • I.1Experimental evidence
    • I.2Species differences in procedural metacognitive skills
  • II.Implicit metacommunication in nonhumans
    • II.1Can nonhumans control their signalling behavior?
      • II.1.1Ostensive communication as a cue for intentional communication
      • II.1.2Re-defining communicative actions
      • II.1.3Operationalizing the presence of first-order intentionality in communication
    • II.2Medium-dependent efficiency: Evolutionary preconditions for individual control and monitoring processes
    • II.3Functional components of procedural metacommunication in nonhumans
      • II.3.1A functional duality
      • II.3.2The puzzle of communicative deception
      • II.3.3Selecting and monitoring informational goals
        • II.3.3.1Metalearning
        • II.3.3.2Informativeness versus complexity (effortfulness)
        • II.3.3.3Comparative urgency
        • II.3.3.4Accuracy versus deception
        • II.3.3.5Clarity as ease of processing
  • III.From nonhuman to human informational communication: The evolution of metacommunication
    • III.1From procedural to explicit metacommunication
    • III.2Discontinuist view: A metacommunicative module
    • III.3A continuist proposal: Metacognitive processes
    • III.4How do continuists explain conversational flexibility?
  • Concluding speculations
  • Acknowledgments
  • Notes
  • References
Published online: 7 July 2023
DOI logo
https://doi.org/10.1075/elt.00046.pro
References (167)
References
Abbot-Smith, K., Schulze, C., Anagnostopoulou, N., Zajączkowska, M., & Matthews, D. (2022). How do 3-year-olds use relevance inferencing to interpret indirect speech?. First Language, 42 (1), 3–21. DOI logoGoogle Scholar
Ackerman, R. (2014). The diminishing criterion model for metacognitive regulation of time investment. Journal of experimental psychology: General, 143 (3), 1349–1368. DOI logoGoogle Scholar
Aguirre, M., Brun, M., Reboul, A., & Mascaro, O. (2022). How do we interpret questions? Simplified representations of knowledge guide humans’ interpretation of information requests. Cognition, 218 1, 104954. DOI logoGoogle Scholar
Akçay, Ç. L., Tom, M. E., Campbell, S. E., & Beecher, M. D. (2013). Song type matching is an honest early threat signal in a hierarchical animal communication system. Proceedings of the Royal Society B: Biological Sciences, 280 (1756), 20122517. DOI logoGoogle Scholar
Allen, J. A., Garland, E. C., Dunlop, R. A., & Noad, M. J. (2018). Cultural revolutions reduce complexity in the songs of humpback whales. Proceedings of the Royal Society B, 285 (1891), 20182088. DOI logoGoogle Scholar
Arnold, K., & Zuberbühler, K. (2012). Call combinations in monkeys: compositional or idiomatic expressions?. Brain and language, 120 (3), 303–309. DOI logoGoogle Scholar
Avital, E., & Jablonka, E. (2000). Animal traditions: Behavioural inheritance in evolution. Cambridge: Cambridge University Press. DOI logoGoogle Scholar
Balcomb, F. K., & Gerken, L. (2008). Three-year-old children can access their own memory to guide responses on a visual matching task. Developmental science, 11 (5), 750–760. DOI logoGoogle Scholar
Barbas, H. (2007). Flow of information for emotions through temporal and orbitofrontal pathways. Journal of anatomy, 211 (2), 237–249. DOI logoGoogle Scholar
Barr, D. J., & Keyser, B. (2002). Anchoring comprehension in linguistic precedents. Journal of Memory and Language, 46 (2), 391–418. DOI logoGoogle Scholar
Basile, B. M., Hampton, R. R., Suomi, S. J., & Murray, E. A. (2009). An assessment of memory awareness in tufted capuchin monkeys (Cebus apella). Animal Cognition, 12 (1), 169–180. DOI logoGoogle Scholar
Beran, M. J., French, K., Smith, T. R., & Parrish, A. E. (2019). Limited evidence of number–space mapping in rhesus monkeys (Macaca mulatta) and capuchin monkeys (Sapajus apella). Journal of Comparative Psychology, 133 (3), 281–293. DOI logoGoogle Scholar
Beran, M. J., & Smith, J. D. (2011). Information seeking by rhesus monkeys (Macaca mulatta) and capuchin monkeys (Cebus apella). Cognition, 120 (1), 90–105. DOI logoGoogle Scholar
Bernard, S., Proust, J., & Clément, F. (2015). Procedural metacognition and false belief understanding in 3-to 5-year-old children. PloS one, 10 (10), e0141321. DOI logoGoogle Scholar
Berwick, R. C., Beckers, G. J., Okanoya, K., & Bolhuis, J. J. (2012). A bird’s eye view of human language evolution. Frontiers in evolutionary neuroscience, 4 1, 5. DOI logoGoogle Scholar
Brown, C., Garwood, M. P., & Williamson, J. E. (2012). It pays to cheat: tactical deception in a cephalopod social signalling system. Biology letters, 8 (5), 729–732. DOI logoGoogle Scholar
Browning, H., & Birch, J. (2022). Animal sentience. Philosophy Compass, 17 (5), e12822. DOI logoGoogle Scholar
Brumm, H., & Slabbekoorn, H. (2005). Acoustic communication in noise. Advances in the Study of Behavior 35 1, 151–209. DOI logoGoogle Scholar
Burkart, J., Martins, E. G., Miss, F., & Zürcher, Y. (2018). From sharing food to sharing information: cooperative breeding and language evolution. Interaction Studies, 19 (1–2), 136–150. DOI logoGoogle Scholar
Byrne, R. W., Cartmill, E., Genty, E., Graham, K. E., Hobaiter, C., & Tanner, J. (2017). Great ape gestures: intentional communication with a rich set of innate signals. Animal cognition, 20 (4), 755–769. DOI logoGoogle Scholar
Call, J. (2012). Seeking information in non-human animals: weaving a metacognitive web. In M. Beran, J. Brandl, J. Perner & J. Proust (eds.), Foundations of metacognition, Oxford: Oxford University press, 62–75. DOI logoGoogle Scholar
Chalcraft, V. J., & Gardner, R. A. (2005). Cross-fostered chimpanzees modulate signs of American Sign Language. Gesture, 5 (1–2), 107–132. DOI logoGoogle Scholar
Cheney, D. L., & Seyfarth, R. M. (1990). How monkeys see the world: Inside the mind of another species. Chicago: University of Chicago Press. DOI logoGoogle Scholar
Cisek, P. (2007). Cortical mechanisms of action selection: the affordance competition hypothesis. Philosophical Transactions of the Royal Society B: Biological Sciences, 362 (1485), 1585–1599. DOI logoGoogle Scholar
(2022). Evolution of behavioural control from chordates to primates. Philosophical Transactions of the Royal Society B, 377 (1844), 20200522. DOI logoGoogle Scholar
Converse, P. D., & DeShon, R. P. (2009). A tale of two tasks: reversing the self-regulatory resource depletion effect. Journal of Applied Psychology, 94 (5), 1318.1324. DOI logoGoogle Scholar
Cunningham, E., & Janson, C. (2007). Integrating information about location and value of resources by white-faced saki monkeys (Pithecia pithecia). Animal cognition, 10 (3), 293–304. DOI logoGoogle Scholar
Daniel, R., & Pollmann, S. (2012). Striatal activations signal prediction errors on confidence in the absence of external feedback. Neuroimage, 59 (4), 3457–3467. DOI logoGoogle Scholar
Dawkins, R., & Krebs, J. R. (1978). Animal signals: Information or manipulation? In J. R. Krebs & N. B. Davies (Eds.), Behavioural ecology: An evolutionary approach (Vol. 21, pp. 282–309). Oxford: Blackwell Scientific Publications.Google Scholar
Deacon, T. W. (2010). A role for relaxed selection in the evolution of the language capacity. Proceedings of the National Academy of Sciences, 107 (supplement_2), 9000–9006. DOI logoGoogle Scholar
de Framond, L., Brumm, H., Thompson, W. I., Drabing, S. M., & Francis, C. D. (2022). The broken-wing display across birds and the conditions for its evolution. Proceedings of the Royal Society B, 289 1(1971), 20220058. DOI logoGoogle Scholar
Dretske, F. (1981). Knowledge and the Flow of Information. Cambridge: Cambridge University PressGoogle Scholar
(1988). Explaining Behavior. Cambridge,Mass: MIT Press.Google Scholar
Duffy, A., Latimer, K. W., Goldberg, J. H., Fairhall, A. L., & Gadagkar, V. (2022). Dopamine neurons evaluate natural fluctuations in performance quality. Cell reports, 38 (13), 110574. DOI logoGoogle Scholar
Dunstone, J., & Caldwell, C. A. (2018). Cumulative culture and explicit metacognition: A review of theories, evidence and key predictions. Palgrave Communications, 4 (1). DOI logoGoogle Scholar
Eliades, S. J., & Miller, C. T. (2017). Marmoset vocal communication: behavior and neurobiology. Developmental neurobiology, 77(3), 286–299. DOI logoGoogle Scholar
Ferrigno, S., Kornell, N., & Cantlon, J. F. (2017). A metacognitive illusion in monkeys. Proceedings of the Royal Society B: Biological Sciences, 284 (1862), 20171541. DOI logoGoogle Scholar
Fitch, W. (2017). Empirical approaches to the study of language evolution. Psychonomic bulletin & review, 24.11: 3–33. DOI logoGoogle Scholar
Fitch, W. T., & Friederici, A. D. (2012). Artificial grammar learning meets formal language theory: an overview. Philosophical Transactions of the Royal Society B: Biological Sciences, 367 (1598), 1933–1955. DOI logoGoogle Scholar
Fleming, S. M., Weil, R. S., Nagy, Z., Dolan, R. J., & Rees, G. (2010). Relating introspective accuracy to individual differences in brain structure. Science, 329 (5998), 1541–1543. DOI logoGoogle Scholar
Franke, M., & Jäger, G. (2016). Probabilistic pragmatics, or why Bayes’ rule is probably important for pragmatics. Zeitschrift für sprachwissenschaft, 35 (1), 3–44. DOI logoGoogle Scholar
Frankfurt, H. (1988). The Importance of what we care about, Cambridge: Cambridge University Press. DOI logoGoogle Scholar
Freeberg, T. M., & Lucas, J. R. (2012). Information theoretical approaches to chick-a-dee calls of Carolina chickadees (Poecile carolinensis). Journal of Comparative Psychology, 126 (1), 68–81. DOI logoGoogle Scholar
Gadagkar, V., Puzerey, P. A., Chen, R., Baird-Daniel, E., Farhang, A. R., & Goldberg, J. H. (2016). Dopamine neurons encode performance error in singing birds. Science, 354 (6317), 1278–1282. DOI logoGoogle Scholar
Gadagkar, V., Puzerey, P. A., & Goldberg, J. H. (2019). Dopamine neurons change their tuning according to courtship context in singing birds. bioRxiv, 822817Google Scholar
Gallistel, C. R. (2020). Where meanings arise and how: Building on Shannon’s foundations. Mind & Language, 35 (3), 390–401. DOI logoGoogle Scholar
Gibson, E., Futrell, R., Piantadosi, S. P., Dautriche, I., Mahowald, K., Bergen, L., & Levy, R. (2019). How efficiency shapes human language. Trends in cognitive sciences, 23 (5), 389–407. DOI logoGoogle Scholar
Godfrey-Smith, P. (2017). The evolution of consciousness in phylogenetic context. In K. Andrew & J. Beck (Eds.), The Routledge Handbook of philosophy of animal minds (pp. 216–226). New York: Routledge. DOI logoGoogle Scholar
Gómez, J. C. (1998). Ostensive behavior in great apes: The role of eye contact. In A. E. Russon, K. A. Bard & S. T. Parker (Eds.), Reaching into thought: The minds of the great apes, (pp. 131–151). New York/Cambridge: Cambridge University Press.Google Scholar
Goto, K., & Watanabe, S. (2012). Large-billed crows (Corvus macrorhynchos) have retrospective but not prospective metamemory. Animal Cognition, 15 (1), 27–35. DOI logoGoogle Scholar
Goupil, L., & Kouider, S. (2019). Developing a reflective mind: from core metacognition to explicit self-reflection. Current Directions in Psychological Science, 28 (4), 403–408. DOI logoGoogle Scholar
Goupil, L., & Proust, J. (2023). Curiosity as a metacognitive feeling. Cognition, 231 1, 105325. DOI logoGoogle Scholar
Grice, P. (1989). Studies in the Way of Words. Harvard University Press.Google Scholar
Hage, S. R. (2018). Dual neural network model of speech and language evolution: new insights on flexibility of vocal production systems and involvement of frontal cortex. Current opinion In behavioral sciences, 21 1, 80–87. DOI logoGoogle Scholar
Hage, S. R., & Nieder, A. (2016). Dual neural network model for the evolution of speech and language. Trends in neurosciences, 39 (12), 813–829. DOI logoGoogle Scholar
Haidle, M. N., Bolus, M., Collard, M., Conard, N. J., Garofoli, D., Lombard, M., Nowell, A., Tennie, C. & Whiten, A. (2015). The nature of culture: an eight-grade model for the evolution and expansion of cultural capacities in hominins and other animals. Journal of Anthropological Sciences. Vol. 931 (2015), pp. 43–70.Google Scholar
Hampton, R. R. (2009). Multiple demonstrations of metacognition in nonhumans: Converging evidence or multiple mechanisms? Comparative Cognition and Behavior Reviews, 41, 17–28. DOI logoGoogle Scholar
Harcourt, A. H. (2015). Humankind. New York: Simon and Schuster.Google Scholar
Hauser, M.D. (1996). The evolution of communication. Cambridge, Mass.: MIT Press.Google Scholar
Hauser, M. D. (1997). Minding the behaviour of deception. In Whiten & Byrne (eds.)., Machiavellian intelligence II: Extensions and evaluations, 12 1 (pp. 112–143). New York/Cambridge: Cambridge University Press. DOI logoGoogle Scholar
Hesler, N., Mundry, R., & Dabelsteen, T. (2012). Are there age-related differences in the song repertoire size of Eurasian blackbirds?. Acta ethologica, 15 (2), 203–210. DOI logoGoogle Scholar
Heyes, C., Bang, D., Shea, N., Frith, C. D., & Fleming, S. M. (2020). Knowing ourselves together: The cultural origins of metacognition. Trends in Cognitive Sciences, 24 (5), 349–362. DOI logoGoogle Scholar
Hisey, E., Kearney, M. G., & Mooney, R. (2018). A common neural circuit mechanism for internally guided and externally reinforced forms of motor learning. Nature neuroscience, 21 (4), 589–597. DOI logoGoogle Scholar
Hobaiter, C., & Byrne, R. W. (2017). What is a gesture? A meaning-based approach to defining gestural repertoires. Neuroscience & Biobehavioral Reviews, 82 1, 3–12. DOI logoGoogle Scholar
Hopkins, W. D., Taglialatela, J. P., & Leavens, D. A. (2007). Chimpanzees differentially produce novel vocalizations to capture the attention of a human. Animal behaviour, 73 (2), 281–286. DOI logoGoogle Scholar
Inzlicht, M., Bartholow, B. D., & Hirsh, J. B. (2015). Emotional foundations of cognitive control. Trends in cognitive sciences, 19 (3), 126–132. DOI logoGoogle Scholar
Jarvis, E. D. (2013). Evolution of brain pathways for vocal learning in birds and humans. In R. C. Berwick & N. Chomsky (Eds.), Birdsong, speech, and language: exploring the evolution of mind and brain (pp. 63–107). Cambridge, Mass.: The MIT Press. DOI logoGoogle Scholar
Kang, M. J., Hsu, M., Krajbich, I. M., Loewenstein, G., McClure, S. M., Wang, J. T. Y., & Camerer, C. F. (2009). The wick in the candle of learning: Epistemic curiosity activates reward circuitry and enhances memory. Psychological Science, 20 (8), 963–973. DOI logoGoogle Scholar
Kepecs, A., & Mainen, Z. F. (2012). A computational framework for the study of confidence in humans and animals. Philosophical Transactions of the Royal Society B: Biological Sciences, 367 (1594), 1322–1337. DOI logoGoogle Scholar
Kershenbaum, A., Demartsev, V., Gammon, D. E., Geffen, E., Gustison, M. L., Ilany, A., & Lameira, A. R. (2021). Shannon entropy as a robust estimator of Zipf’s Law in animal vocal communication repertoires. Methods in Ecology and Evolution, 12 (3), 553–564. DOI logoGoogle Scholar
Keysar, B., Lin, S., & Barr, D. J. (2003). Limits on theory of mind use in adults. Cognition, 89 (1), 25–41. DOI logoGoogle Scholar
Kissine, M. (2016). Pragmatics as metacognitive control. Frontiers in psychology, 6 1, 2057. DOI logoGoogle Scholar
Koechlin, E. (2014). An evolutionary computational theory of prefrontal executive function in decision-making. Philosophical Transactions of the Royal Society B: Biological Sciences, 369 (1655), 20130474. DOI logoGoogle Scholar
Koechlin, E., & Summerfield, C. (2007). An information theoretical approach to prefrontal executive function. Trends in cognitive sciences, 11 (6), 229–235. DOI logoGoogle Scholar
Konishi, M. (2004). The role of auditory feedback in birdsong. Annals of the New York Academy of Sciences, 1016 (1), 463–475. DOI logoGoogle Scholar
Koriat, A. (2000). The feeling of knowing: Some metatheoretical implications for consciousness and control. Consciousness and cognition, 9 (2), 149–171. DOI logoGoogle Scholar
(2008). Subjective confidence in one’s answers: the consensuality principle. Journal of Experimental Psychology: Learning, Memory, and Cognition, 34 (4), 945–959.Google Scholar
Koriat, A., & Ackerman, R. (2010). Metacognition and mindreading: Judgments of learning for self and other during self-paced study. Consciousness and cognition, 19 (1), 251–264. DOI logoGoogle Scholar
Koriat, A., & Levy-Sadot, R. (1999). Processes underlying metacognitive judgments: Information-based and experience-based monitoring of one’s own knowledge. In S. Chaïken & Y. Trope (Eds.), Dual Process theories in social psychology (pp. 483–502). New York: Guilford.Google Scholar
Koriat, A., Ma’ayan, H., & Nussinson, R. (2006). The intricate relationships between monitoring and control in metacognition: lessons for the cause-and-effect relation between subjective experience and behavior. Journal of experimental psychology: general, 135 (1), 36–69. DOI logoGoogle Scholar
Koriat, A., Nussinson, R., Bless, H., & Shaked, N. (2008). Information-based and experience-based metacognitive judgments: Evidence from subjective confidence. In J. Dunloski and R. A. Bjork, (eds.) A Handbook of Memory and Metamemory, Psychology Press, 117–136.Google Scholar
Kornell, N., Son, L. K., & Terrace, H. S. (2007). Transfer of metacognitive skills and hint seeking in monkeys. Psychological Science, 18 (1), 64–71. DOI logoGoogle Scholar
Kuchling, F., Fields, C., & Levin, M. (2022). Metacognition as a consequence of competing evolutionary time scales. Entropy, 24 (5), 601. DOI logoGoogle Scholar
Kurzban, R., Duckworth, A., Kable, J. W., & Myers, J. (2013). An opportunity cost model of subjective effort and task performance. Behavioral and brain sciences, 36 (6), 661–679. DOI logoGoogle Scholar
Lage, C. A., Wolmarans, D. W., & Mograbi, D. C. (2022). An evolutionary view of self-awareness. Behavioural Processes, 194 1, 104543. DOI logoGoogle Scholar
Lak, A., Stauffer, W. R., & Schultz, W. (2014). Dopamine prediction error responses integrate subjective value from different reward dimensions. Proceedings of the National Academy of Sciences, 111 (6), 2343–2348. DOI logoGoogle Scholar
Lambert, M. L., & Osvath, M. (2020). Investigating information seeking in ravens (Corvus corax). Animal Cognition, 23 (4), 671–680. DOI logoGoogle Scholar
Lameira, A. R. (2017). Bidding evidence for primate vocal learning and the cultural substrates for speech evolution. Neuroscience & Biobehavioral Reviews, 83 1, 429–439. DOI logoGoogle Scholar
Liszkowski, U., Carpenter, M., & Tomasello, M. (2008). Twelve-month-olds communicate helpfully and appropriately for knowledgeable and ignorant partners. Cognition, 108 (3), 732–739. DOI logoGoogle Scholar
Macedonia, J. M., & Evans, C. S. (1993). Essay on contemporary issues in ethology: Variation among mammalian alarm call systems and the problem of meaning in animal signals. Ethology, 93 (3), 177–197. DOI logoGoogle Scholar
Magrath, R. D., Haff, T. M., Horn, A. G., & Leonard, M. L. (2010). Calling in the face of danger. Predation risk and acoustic communication by parent birds and their offspring. In J. C. Mitani, H. J. Brockman, T. J. Roper, M. Naguib & K. E. Wynne-Edwards (Eds.), Advances in the Study of Behavior (Vol. 411, pp. 187–253). New York: Academic Press.Google Scholar
Margoliash, D., & Nusbaum, H. C. (2009). Language: the perspective from organismal biology. Trends in cognitive sciences, 13 (12), 505–510. DOI logoGoogle Scholar
Marler, P. (1967). Animal Communication Signals: We are beginning to understand how the structure of animal signals relates to the function they serve. Science, 157 (3790), 769–774. DOI logoGoogle Scholar
Marler, P., Karakashian, S., & Gyger, M. (1991). Do animals have the option of withholding signals when communication is inappropriate? The audience effect. In C. A. Ristau (Ed.), Cognitive ethology: The minds of other animals: Essays in honor of Donald R. Griffin (pp. 187–208). Mawah: Lawrence Erlbaum Associates, Inc.Google Scholar
Maynard Smith, J. & Harper, D. (2003). Animal signals. Oxford: Oxford University Press.Google Scholar
Middlebrooks, P. G., & Sommer, M. A. (2012). Neuronal correlates of metacognition in primate frontal cortex. Neuron, 75 (3), 517–530. DOI logoGoogle Scholar
Miller, C. T. & Thomas, A. W. (2012). Individual recognition during bouts of antiphonal calling in common marmosets. Journal of Comparative Physiology, 1981: 337–346. DOI logoGoogle Scholar
Millikan, R. (1984) Language, Thought and Other Biological Categories , Cambridge, Mass:MIT press (ISBN 978-0262631150) DOI logoGoogle Scholar
Millikan, R. G. (2005). Language: A biological model. New York: Oxford University Press. DOI logoGoogle Scholar
Millikan, R. G. (2017). Beyond concepts: Unicepts, language, and natural information. New York/Oxford: Oxford University Press. DOI logoGoogle Scholar
Mitani, J. C., & Watts, D. P. (2005). Correlates of territorial boundary patrol behaviour in wild chimpanzees. Animal Behaviour, 70 (5), 1079–1086. DOI logoGoogle Scholar
Moore, R. (2016). Meaning and ostension in great ape gestural communication. Animal Cognition, 19 (1), 223–231. DOI logoGoogle Scholar
Morales, J., Lau, H., & Fleming, S. M. (2018). Domain-general and domain-specific patterns of activity supporting metacognition in human prefrontal cortex. Journal of Neuroscience, 38 (14), 3534–3546. DOI logoGoogle Scholar
Nowicki, S., & Searcy, W. A. (2005). Song and mate choice in birds: how the development of behavior helps us understand function. The Auk, 122(1), 1–14. DOI logoGoogle Scholar
Ouattara, K., Lemasson, A., & Zuberbühler, K. (2009). Campbell’s monkeys use affixation to alter call meaning. PloS one, 4 (11), e7808. DOI logoGoogle Scholar
Oudeyer, P. Y., & Smith, L. B. (2016). How evolution may work through curiosity-driven developmental process. Topics in Cognitive Science, 8 (2), 492–502. DOI logoGoogle Scholar
Paulus, M., Proust, J., & Sodian, B. (2013). Examining implicit metacognition in 3.5-year-old children: An eye-tracking and pupillometric study. Frontiers in Psychology, 4 1, 145. DOI logoGoogle Scholar
Petkov, C. I., & Jarvis, E. D. (2012). Birds, primates, and spoken language origins: behavioral phenotypes and neurobiological substrates. Frontiers in evolutionary neuroscience, 4 1, 12. DOI logoGoogle Scholar
Pezzulo, G., & Cisek, P. (2016). Navigating the affordance landscape: feedback control as a process model of behavior and cognition. Trends in cognitive sciences, 20 (6), 414–424. DOI logoGoogle Scholar
Pinto, A., Oates, J., Grutter, A., & Bshary, R. (2011). Cleaner wrasses Labroides dimidiatus are more cooperative in the presence of an audience. Current Biology, 21 (13), 1140–1144. DOI logoGoogle Scholar
Planer, R. (2017a). Talking About Tools: Did Early Pleistocene Hominins Have a Protolanguage? Biological Theory, 121, 211–221. DOI logoGoogle Scholar
(2017b). Protolanguage might have evolved before ostensive communication. Biological Theory, 12 1, 72–84. DOI logoGoogle Scholar
Poole, J. H., Tyack, P. L., Stoeger-Horwath, A. S., & Watwood, S. (2005). Elephants are capable of vocal learning. Nature, 434 (7032), 455–456. DOI logoGoogle Scholar
Proust, J. (2012). Metacognition and mindreading: one or two functions? in: M. Beran, J. Brandl, J. Perner & J. Proust (Eds.), The Foundations of Metacognition (pp. 234–251). Oxford: Oxford: Oxford University Press. DOI logoGoogle Scholar
(2013). The Philosophy of Metacognition. Oxford: Oxford University Press. DOI logoGoogle Scholar
(2015). Time and action: Impulsivity, habit, strategy. Review of Philosophy and Psychology, 6 (4), 717–743. DOI logoGoogle Scholar
(2016). The evolution of primate communication and metacommunication. Mind & Language, 31 (2), 177–203. DOI logoGoogle Scholar
(2019). From comparative studies to interdisciplinary research on metacognition. Animal Behavior and Cognition, 6 (4), 309–328. DOI logoGoogle Scholar
(2022). The cultural evolution of information seeking, Journal of Cognition and Culture, 22 (5), 467–484.Google Scholar
(2023). A semantics for affordances, Philosophical Psychology.Google Scholar
Pulvermüller, F., & Grisoni, L. (2020). Semantic prediction in brain and mind. Trends in Cognitive Sciences, 24 (10), 781–784. DOI logoGoogle Scholar
Reboul, A. (2011). A relevance-theoretic account of the evolution of implicit communication. Studies in Pragmatics, 13 (1), 1–19.Google Scholar
Rendall, D., Owren, M. J., & Ryan, M. J. (2009). What do animal signals mean?. Animal Behaviour, 78 (2), 233–240. DOI logoGoogle Scholar
Rhodes, M. G., & Castel, A. D. (2009). Metacognitive illusions for auditory information: Effects on monitoring and control. Psychonomic Bulletin & Review, 16 (3), 550–554. DOI logoGoogle Scholar
Roberts, C. (2012). Information Structure: Towards an integrated formal theory of pragmatics. Semantics and Pragmatics, 5 (0), 61:1–69. DOI logoGoogle Scholar
Roberts, W. A., Feeney, M. C., McMillan, N., MacPherson, K., Musolino, E., & Petter, M. (2009). Do pigeons (Columba livia) study for a test? Journal of Experimental Psychology: Animal Behavior Processes, 35 1, 129–142.Google Scholar
Roberts, W. A., McMillan, N., Musolino, E., & Cole, M. (2012). Information seeking in animals: metacognition?. Comparative Cognition & Behavior Reviews, 7 1, 85–109. DOI logoGoogle Scholar
Rosati, A. G., & Santos, L. R. (2016). Spontaneous metacognition in rhesus monkeys. Psychological Science, 27 (9), 1181–1191. DOI logoGoogle Scholar
Salmi, R., Szczupider, M., & Carrigan, J. (2022). A novel attention-getting vocalization in zoo-housed western gorillas. Plos one, 17 (8), e0271871. DOI logoGoogle Scholar
Santi, A., Adams, A., & Bassett, J. (2010). The role of keypecking during filled intervals on the judgment of time for empty and filled intervals by pigeons. Learning & Behavior, 38 (1), 42–49. DOI logoGoogle Scholar
Scarantino, A., & Clay, Z. (2015). Contextually variable signals can be functionally referential. Animal Behaviour, 100(100), e1–e8. DOI logoGoogle Scholar
Schel, A. M., Candiotti, A., & Zuberbühler, K. (2010). Predator-deterring alarm call sequences in Guereza colobus monkeys are meaningful to conspecifics. Animal Behaviour, 80 (5), 799–808. DOI logoGoogle Scholar
Schlenker, P. (2018). Iconic pragmatics. Natural Language & Linguistic Theory, 36 (3), 877–936. DOI logoGoogle Scholar
Schlenker, P., Chemla, E., & Zuberbühler, K. (2016). What do monkey calls mean?. Trends in Cognitive Sciences, 20 (12), 894–904. DOI logoGoogle Scholar
Schwartz, B. L. (2019). Using natural ecology to predict higher cognition in human and non-human primates. Animal Behavior and Cognition, 6 (4), 344–354. DOI logoGoogle Scholar
Schwarz, N., & Clore, G. L. (2007). Feelings and phenomenal experiences. In A. W. Kruglanski, & Higgins, (Eds.). (2013). Social psychology: Handbook of basic principles (pp. 385–407). New York: Guilford.Google Scholar
Scott-Phillips, T. C. (2016). Meaning in great ape communication: summarising the debate. Animal Cognition, 19 (1), 233–238. DOI logoGoogle Scholar
Seyfarth, R. M., & Cheney, D. L. (2003). Signallers and recipients in animal communication. Annual review of psychology, 54 (1), 145–173. DOI logoGoogle Scholar
Shea, N., Boldt, A., Bang, D., Yeung, N., Heyes, C., & Frith, C. D. (2014). Supra-personal cognitive control and metacognition. Trends in cognitive sciences, 18 (4), 186–193. DOI logoGoogle Scholar
Shea, N., Godfrey-Smith, P., & Cao, R. (2018). Content in simple signalling systems. The British Journal for the Philosophy of Science, 691, 4. DOI logoGoogle Scholar
Sherman, P. W. (1977). Nepotism and the Evolution of Alarm Calls: Alarm calls of Belding’s ground squirrels warn relatives, and thus are expressions of nepotism. Science, 197 (4310), 1246–1253. DOI logoGoogle Scholar
Skyrms, B. (2010). Signals: Evolution, Learning, and Information. Oxford: Oxford University Press. DOI logoGoogle Scholar
Slocombe, K. E., & Zuberbühler, K. (2007). Chimpanzees modify recruitment screams as a function of audience composition. Proceedings of the National Academy of Sciences, 104 (43), 17228–17233. DOI logoGoogle Scholar
Sperber, D. (2000). Metarepresentations in an evolutionary perspective. In D. Sperber (ed.), Metarepresentations: A multidisciplinary perspective (pp. 117–137). Oxford: Oxford University Press.Google Scholar
(2019). Personal notes on a shared trajectory. In K. Scott, B. Clark & R. Carston (Eds.), Relevance, Pragmatics and Interpretation (pp. 13–20). Cambridge: Cambridge University Press. DOI logoGoogle Scholar
Sperber, D., & Wilson, D. (1986/1995). Relevance: Communication and cognition (Vol. 1421). Cambridge, MA: Harvard University Press.Google Scholar
(2002). Pragmatics, modularity and mind-reading. Mind & language, 17 (1–2), 3–23. DOI logoGoogle Scholar
Sterelny, K. (2016). Cumulative cultural evolution and the origins of language. Biological Theory, 11 (3), 173–186. DOI logoGoogle Scholar
Sutton, J. E., & Shettleworth, S. J. (2008). Memory without awareness: pigeons do not show metamemory in delayed matching to sample. Journal of Experimental Psychology: Animal Behavior Processes, 34 (2), 266–282.Google Scholar
Suzuki, T. N. (2016). Semantic communication in birds: evidence from field research over the past two decades. Ecological Research, 31 (3), 307–319. DOI logoGoogle Scholar
Takahashi, D. Y., Fenley, A. R., Teramoto, Y., Narayanan, D. Z., Borjon, J. I., Holmes, P., & Ghazanfar, A. A. (2015). The developmental dynamics of marmoset monkey vocal production. Science, 349 (6249), 734–738. DOI logoGoogle Scholar
Tang, H., Costa, V. D., Bartolo, R., and Averbeck, B. B. (2022). Differential coding of goals and actions in ventral and dorsal corticostriatal circuits during goal-directed behavior. Cell Reports 38 1, 110198. DOI logoGoogle Scholar
Terrace, H. S., & Son, L. K. (2009). Comparative metacognition. Current opinion in neurobiology, 19 (1), 67–74. DOI logoGoogle Scholar
Tian, J., Huang, R., Cohen, J. Y., Osakada, F., Kobak, D., Machens, C. K., & Watabe-Uchida, M. (2016). Distributed and mixed information in monosynaptic inputs to dopamine neurons. Neuron, 91 (6), 1374–1389. DOI logoGoogle Scholar
Townsend, S. W., Koski, S. E., Byrne, R. W., Slocombe, K. E., Bickel, B., Boeckle, M., … & Manser, M. B. (2017). Exorcising Grice’s ghost: An empirical approach to studying intentional communication in animals. Biological Reviews, 92 (3), 1427–1433. DOI logoGoogle Scholar
Tsujimoto, S., Genovesio, A., & Wise, S. P. (2010). Evaluating self-generated decisions in frontal pole cortex of monkeys. Nature neuroscience, 13 (1), 120–126. DOI logoGoogle Scholar
van Schaik, C. P., Pradhan, G. R., & Tennie, C. (2019). Teaching and curiosity: sequential drivers of cumulative cultural evolution in the hominin lineage. Behavioral ecology and sociobiology, 73 (1), 1–11. DOI logoGoogle Scholar
Watanabe, A., & Clayton, N. S. (2016). Hint-seeking behaviour of western scrub-jays in a metacognition task. Animal cognition, 19 (1), 53–64. DOI logoGoogle Scholar
Wheeler, B. C., & Fischer, J. (2012). Functionally referential signals: a promising paradigm whose time has passed. Evolutionary Anthropology: Issues, News, and Reviews, 21 (5), 195–205. DOI logoGoogle Scholar
Whiten, A., & Byrne, R. W. (1988). Tactical deception in primates. Behavioral and brain sciences, 11 (2), 233–244. DOI logoGoogle Scholar
(Eds.). (1997). Machiavellian intelligence II: Extensions and evaluations. Cambridge, Cambridge University Press. DOI logoGoogle Scholar
Wich, S. A., Krützen, M., Lameira, A. R., Nater, A., Arora, N., Bastian, M. L., … & van Schaik, C. P. (2012). Call cultures in orang-utans?. PLoS one, 7 (5), e36180. DOI logoGoogle Scholar
Wismer, S., Pinto, A. I., Vail, A. L., Grutter, A. S., & Bshary, R. (2014). Variation in cleaner wrasse cooperation and cognition: influence of the developmental environment? Ethology, 120 (6), 519–531. DOI logoGoogle Scholar
Wu, S., Barr, D. J., Gann, T. M., & Keysar, B. (2013). How culture influences perspective taking: differences in correction, not integration. Frontiers in human neuroscience, 7 1, 822. DOI logoGoogle Scholar
Xiao, L., Chattree, G., Oscos, F. G., Cao, M., Wanat, M. J., & Roberts, T. F. (2018). A basal ganglia circuit sufficient to guide birdsong learning. Neuron, 98 (1), 208–221. DOI logoGoogle Scholar
Zakrzewski, A. C., Perdue, B. M., Beran, M. J., Church, B. A., & Smith, J. D. (2014). Cashing out: The decisional flexibility of uncertainty responses in rhesus macaques (Macaca mulatta) and humans (Homo sapiens). Journal of Experimental Psychology: Animal Learning and Cognition, 40 (4), 490–501.Google Scholar
Zuberbühler, K. (2008). Audience effects. Current Biology, 18 (5), 189–190. DOI logoGoogle Scholar