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This work was supported by the European Union's Seventh Framework Programme (FP7/2007–2013)/ERC Grant 609819 (SOMICS). ; Communication, when defined as an act intended to affect the psychological state of another individual, demands the use of inference. Either the signaler, the recipient, or both must make leaps of understanding which surpass the semantic information available and draw from pragmatic clues to fully imbue and interpret meaning. While research into human communication and the evolution of language has long been comfortable with mentalistic interpretations of communicative exchanges, including rich attributions of mental state, research into animal communication has balked at theoretical models which describe mentalized cognitive mechanisms. We submit a new theoretical perspective on animal communication: the model of inferential communication. For use when existing proximate models of animal communication are not sufficient to fully explain the complex, flexible, and intentional communication documented in certain species, specifically non-human primates, we present our model as a bridge between shallower, less cognitive descriptions of communicative behavior and the perhaps otherwise inaccessible mentalistic interpretations of communication found in theoretical considerations of human language. Inferential communication is a framework that builds on existing evidence of referentiality, intentionality, and social inference in primates. It allows that they might be capable of applying social inferences to a communicative setting, which could explain some of the cognitive processes that enable the complexity and flexibility of primate communication systems. While historical models of animal communication focus on the means-ends process of behavior and apparent cognitive outcomes, inferential communication invites consideration of the mentalistic processes that must underlie those outcomes. We propose a mentalized approach to questions, investigations, and interpretations of non-human primate communication. We include an overview of both ultimate and proximate models of animal communication, which contextualize the role and utility of our inferential communication model, and provide a detailed breakdown of the possible levels of cognitive complexity which could be investigated using this framework. Finally, we present some possible applications of inferential communication in the field of non-human primate communication and highlight the role it could play in advancing progress toward an increasingly precise understanding of the cognitive capabilities of our closest living relatives. ; Publisher PDF ; Peer reviewed

AbstractWe investigated whether great apes, like human infants, monkeys and dogs, are subject to a strong gravity bias when tested with the tubes task, and – in case of mastery – what the source of competence on the tubes task is. We presented 22 apes with three versions of the tubes task, in which an object is dropped down a tube connected to one of three potential hiding places and the subject is required to locate the object. In two versions, apes were confronted with a causal tube that varied in the amount of perceptual information it provided (i.e. presence or absence of acoustic cues). The third version was a non‐causal adaptation of the task in which a painted line 'connected' dropping and hiding places. Results indicate that apes neither have a reliable gravity bias when tested with the tubes, nor understand the causal function of the tube. Even though there is evidence that they can integrate tube‐related causal information to localize the object, they seem to depend mainly on non‐causal inferences when searching for an invisibly displaced object.

The study was funded by the European Union's Horizon 2020 Research and Innovation Program under the Marie Skłodowska-Curie grant agreement no. 702137 attributed to A.R.L. ; One of the defining features of language is displaced reference—the capacity to transmit information about something that is not present or about a past or future event. It is very rare in nature and has not been shown in any nonhuman primate, confounding, as such, any understanding of its precursors and evolution in the human lineage. Here, we describe a vocal phenomenon in a wild great ape with unparalleled affinities with displaced reference. When exposed to predator models, Sumatran orangutan mothers temporarily suppressed alarm calls up to 20 min until the model was out of sight. Subjects delayed their vocal responses in function of perceived danger for themselves, but four major predictions for stress-based mechanisms were not met. Conversely, vocal delay was also a function of perceived danger for another—an infant—suggesting high-order cognition. Our findings suggest that displaced reference in language is likely to have originally piggybacked on akin behaviors in an ancestral hominid. ; Publisher PDF ; Peer reviewed

Abstract
Pragmatic theories assume that during communicative exchanges humans strive to be optimally informative and spontaneously adjust their communicative signals to satisfy their addressee's inferred epistemic needs. For instance, when necessary, adults flexibly and appropriately modify their communicative gestures to provide their partner the relevant information she lacks about the situation. To investigate this ability in infants, we designed a cooperative task in which 18-month-olds were asked to point at the target object they wanted to receive. In Experiment 1, we found that when their desired object was placed behind a distractor object, infants appropriately modified their prototypical pointing to avoid mistakenly indicating the distractor to their partner. When the objects were covered, and their cooperative partner had no information (Experiment 2) or incorrect information (Experiment 3) about the target's location – as opposed to being knowledgeable about it – infants pointed differentially more often at the target and employed modified pointing gestures more frequently as a function of the amount of relevant information that their partner needed to retrieve their desired object from its correct location. These findings demonstrate that when responding to a verbal request in a cooperative task 18-month-old infants can take into account their communicative partner's epistemic states and when necessary provide her with the relevant information she lacks through sufficiently informative deictic gestures. Our results indicate that infants possess an early emerging, species-unique cognitive adaptation specialized for communicative mindreading and pragmatic inferential communication which enable the efficient exchange of relevant information between communicating social partners in cooperative contexts.

This project has received funding from the European Research Council (ERC) under the European Union's Horizon 2020 Research and Innovation Programme (Grant Agreement No. 639072). ; Human adults can infer unseen causes because they represent the events around them in terms of their underlying causal mechanisms. It has been argued that young preschoolers can also make causal inferences from an early age, but whether or not nonhuman apes can go beyond associative learning when exploiting causality is controversial. However, much of the developmental research to date has focused on fully-perceivable causal relations or highlighted the existence of a causal relationship verbally and these were found to scaffold young children's abilities. We examined inferences about unseen causes in children and chimpanzees in the absence of linguistic cues. Children (N=129, aged 3-6 years) and zoo-living chimpanzees (N=11, aged 7-41 years) were presented with an event in which a reward was dropped through an opaque forked-tube into one of two cups. An auditory cue signaled which of the cups contained the reward. In the causal condition, the cue followed the dropping event, making it plausible that the sound was caused by the reward falling into the cup; and in the arbitrary condition, the cue preceded the dropping event, making the relation arbitrary. By 4-years of age, children performed better in the causal condition than the arbitrary one, suggesting that they engaged in reasoning. A follow-up experiment ruled out a simpler associative learning explanation. Chimpanzees and 3-year-olds performed at chance in both conditions. These groups' performance did not improve in a simplified version of the task involving shaken boxes; however, the use of causal language helped 3-year-olds. The failure of chimpanzees could reflect limitations in reasoning about unseen causes or a more general difficulty with auditory discrimination learning. ; Publisher PDF ; Peer reviewed

AbstractThe recognition of iconic correspondence between signal and referent has been argued to bootstrap the acquisition and emergence of language. Here, we study the ontogeny, and to some extent the phylogeny, of the ability to spontaneously relate iconic signals, gestures, and/or vocalizations, to previous experience. Children at 18, 24, and 36 months of age (N = 216) and great apes (N = 13) interacted with two apparatuses, each comprising a distinct action and sound. Subsequently, an experimenter mimicked either the action, the sound, or both in combination to refer to one of the apparatuses. Experiments 1 and 2 found no spontaneous comprehension in great apes and in 18‐month‐old children. At 24 months of age, children were successful with a composite vocalization‐gesture signal but not with either vocalization or gesture alone. At 36 months, children succeeded both with a composite vocalization‐gesture signal and with gesture alone, but not with vocalization alone. In general, gestures were understood better compared to vocalizations. Experiment 4 showed that gestures were understood irrespective of how children learned about the corresponding action (through observation or self‐experience). This pattern of results demonstrates that iconic signals can be a powerful way to establish reference in the absence of language, but they are not trivial for children to comprehend and not all iconic signals are created equal.

Abstract In the context of an imitation game, 12‐ and 18‐month‐old infants saw an adult do such things as make a toy mouse hop across a mat (with sound effects). In one condition (House), the adult ended by placing the mouse in a toy house, whereas in another condition (No House) there was no house present at the final location. Infants at both ages usually simply put the mouse in the house (ignoring the hopping motion and sound effects) in the House condition, presumably because they interpreted the adult's action in terms of this final goal and so ignored the behavioral means. In contrast, infants copied the adult's action (both the hopping motion and the sound effects) when no house was present, presumably because here infants saw the action itself as the adult's only goal. From very early, infants' social learning is flexible: infants focus on and copy either the end or the means of an adult action as required by the context.

This research was supported by the European Research Council under the European Union's Seventh Framework Program ( FP7/2007-2013 )/ERC grant agreement no 609819, SOMICS. ; Humans can tell when they find a task difficult. Subtle uncertainty behaviors like changes in motor speed and muscle tension precede and affect these experiences. Theories of animal metacognition likewise stress the importance of endogenous signals of uncertainty as cues that motivate metacognitive behaviors. However, while researchers have investigated second-order behaviors like information seeking and declining difficult trials in nonhuman animals, they have devoted little attention to the behaviors that express the cognitive conflict that gives rise to such behaviors in the first place. Here we explored whether three chimpanzees would, like humans, show hand wavering more when faced with more difficult choices in a touch screen transitive inference task. While accuracy was very high across all conditions, all chimpanzees wavered more frequently in trials that were objectively more difficult, demonstrating a signature behavior which accompanies experiences of difficulty in humans. This lends plausibility to the idea that feelings of uncertainty, like other emotions, can be studied in nonhuman animals. We propose to routinely assess uncertainty behaviors to inform models of procedural metacognition in nonhuman animals. ; Publisher PDF ; Peer reviewed

AbstractBy three years of age, children are skilled at assessing under which circumstances others can see things. However, nothing is known about whether they can use this knowledge to guide their own deceptive behaviour. Here we investigated 3‐year‐olds' ability to strategically inhibit or conceal forbidden actions that a nearby adult experimenter could see or hear. In the first experiment, children were more likely to disobey the adult when she did not have visual access to their activities than they were when she was looking at them. In the second experiment, in which the adult could never see the child, children refrained from making noise when engaging in a prohibited action that the adult might hear. These results suggest that by three years of age children use their knowledge of others' perceptual states to decide whether it is safe to commit a transgression and, moreover, actively conceal perceptual cues that could reveal to others their ongoing transgression.

This research was supported by the European Research Council (ERC) under the European Union's Seventh Framework Programme (FP7/2007–2013)/ERC Grant 609819 (SOMICS). Manuel Bohn was supported by the European Union's Horizon 2020 research and innovation programme under the Marie Sklodowska-Curie grant agreement No 749229. ; Great apes are able to request objects from humans by pointing. It is unclear, however, whether this is an associated response to a certain set of cues (e.g. the presence and attention of a human addressee) or a communicative signal which can be adjusted to relevant aspects of the spatial and social context. In three experiments, we tested captive great apes' flexible use of pointing gestures. We manipulated the communicative context so that the default pointing response of apes would have indicated an undesired object, either due to 1) the spatial arrangements of the target objects, 2) the perspective of the addressee or 3) the knowledge of the addressee about the target objects' location. The results of the three experiments indicate that great apes can successfully adjust their pointing to the spatial configuration of the referent environment such as distance and location of food. However, we found no evidence that they take the perspective or the knowledge of the addressee into account when doing so. This implies that pointing in great apes is a context-sensitive, but maybe less versatile, communicative signal compared to human pointing. ; Publisher PDF ; Peer reviewed

AbstractA key skill in early human development is the ability to comprehend communicative intentions as expressed in both nonlinguistic gestures and language. In the current studies, we confronted domestic dogs (some of whom knew many human 'words') with a task in which they had to infer the intended referent of a human's communicative act via iconic signs – specifically, replicas and photographs. Both trained and untrained dogs successfully used iconic replicas to fetch the desired item, with many doing so from the first trial. Dogs' ability to use photographs in this same situation was less consistent. Because simple matching to sample in experimental contexts typically takes hundreds of trials (and because similarity between iconic sign and target item did not predict success), we propose that dogs' skillful performance in the current task reflects important aspects of the comprehension of human communicative intentions.

AbstractA nonverbal false belief task was administered to children (mean age 5 years) and two great ape species: chimpanzees (Pan troglodytes) and bonobos (Pan paniscus). Because apes typically perform poorly in cooperative contexts, our task was competitive. Two versions were run: in both, a human competitor witnessed an experimenter hide a reward in one of two containers. When the competitor then left the room (version A) or turned around (version B), the experimenter switched the locations of the containers. The competitor returned and reached with effort, but unsuccessfully, towards the incorrect container. Children displayed an understanding of the competitor's false belief by correctly choosing theothercontainer to find the reward. Apes did not. However, in version A (but not version B), apes looked more often at the unchosen container in false belief trials than in true belief control trials, possibly indicating some implicit or uncertain understanding that needs to be investigated further.