Abstract: An important question in the study of animal communication is whether non-human animals are able to produce communicative gestures, i.e. nonvocal bodily actions directed to a recipient, physically ineffective but with a meaning shared in the social group [1]. Passive gestures are instrumental, tuned to the mere presence/absence of others, whereas active informers recognize receivers as communicative agents and activate shared-attention mechanisms for identifying their attentional state (SAM [2]; e.g. Schwab and Huber [3]). Six operational criteria must be evaluated to classify a signal as referential and intentional [4]: (1) alternative gazes between the partner and the target; (2) apparent attention-getting behaviours are deployed; (3) an audience is required to exhibit the behaviour; (4) the attentional status of an observer influences the propensity to exhibit behaviours; (5) communication is persistent and (6) there is elaboration of communicative behaviour when apparent attempts to manipulate the partner fail. Dogs [5] and non-human primates (reviewed in Liebal and Call [6]) can tune a human receiver’s attention to the object of interest by combining directional and attention-getting signals, such as turning the head or body, gazing to the receiver, and/or establishing eye contact. Research on other species is scarce.
Horses rely on humans to survive in domestic settings and may have evolved skills for communicating flexibly with them [7]. Horses understand human attentional cues (such as body and head orientation, eyes opened/closed) [8], permanent pointing [9] and, to some extent, gazing [10]. Here we tested the ability of 14 outdoor, herd-living domestic horses to communicate referentially with a human partner about the location of a desired target, a bucket of food out of reach. After the baiting of two buckets placed in opposite, unreachable locations were shown by the experimenter, the subject would walk to one of the two buckets. Because approaching a bucket would reveal that the food is out of reach, we expected the horse to look back to the experimenter, then to the bucket, and alternate this gazing several times to indicate its intention. To test whether our prediction is correct and alternate gazing is indeed the result of the horse's referential communication, we video-recorded the behaviour of the subjects in the test (FORWARD) and three control conditions: (1) FORWARD: experimenter oriented to the center of the arena, (2) BACK: experimenter backward oriented in respect to the arena, (3) ALONE: experimenter absent, (4) MANY: as FORWARD plus a familiar human oriented to the subject behind the bucket (Figure 1). We used a conservative criterion of back gazing by considering only turning the head back more than 90 degrees. The results confirmed our prediction. The horses alternated gazes between the partner and the buck significantly more often in the FORWARD than in all the other conditions (Table 1), thus satisfying operational criteria #1, #3 and #4. They also alternated head nods with gazes to the partner significantly more often during the FORWARD condition. We thus considered head nods not an instrumental signal of arousal, but an attention-getting behaviour with communicative function. Subjects used both head nods and neck stretched toward the buck more often in the FORWARD than in the BACK and the ALONE conditions, thus satisfying criteria #2, #3 and #4. In condition MANY, the frequency of head nods did not differ from condition FORWARD, probably because nods were directed to the additional partner behind the buck. This also satisfies criteria #4. The horses gazed to the partner most often in the FORWARD than in the BACK and the MANY conditions, but not in the ALONE. In this condition, subjects could observe the partner walking further from the test arena. To test for the different functions of gazes in presence and in absence of the partner, we compared their average duration between the two conditions: the significantly longer duration of gazes when the subject was alone suggests the instrumental monitoring function of gazes in this experimental condition.
Altogether, the findings suggest that domestic horses possess the ability to use referential communication in an interspecific context, but additional analyses are needed to test for operational criteria #5 and #6. Flexible and voluntary use of communicative signals reveal sophisticated cognitive processes involved in the strategic emission of these signals, and the finding of referential communication skills in an ungulate species forces us to reconsider the evolutionary path of intelligence. Furthermore, ungulates are used intensively by humans (transportation, meat, agriculture, leisure activities), and their welfare is often compromised. Determining whether ungulates can communicate their needs and preferences is paramount to a proper ethical management.

Abstract: Both in humans and non-human animals, it has been shown that individuals attend more to those they have previously interacted with and/or they are more closely associated with than to unfamiliar individuals. Whether this preference is mediated by mere social familiarity based on exposure or by the specific relationship between the two individuals, however, remains unclear. The domestic dog is an interesting subject in this line of research as it lives in the human environment and regularly interacts with numerous humans, yet it often has a particularly close relationship with its owner. Therefore, we investigated how long dogs (Canis familiaris) would attend to the actions of two familiar humans and one unfamiliar experimenter, while varying whether dogs had a close relationship with only one or both familiar humans. Our data provide evidence that social familiarity by itself cannot account for dogs’ increased attention towards their owners since they only attended more to those familiar humans with whom they also had a close relationship.

Abstract: Abstract: Understanding animals" abilities to cooperate with and learn from each other has been an active field of research in recent years. One important basis for all types of social interactions is the disposition of animals to pay attention to each other-a factor often neglected in discussions and experiments. Since attention differs between species as well as between individuals, it is likely to influence the amount and type of information different species and/or observers may extract from conspecifics in any given situation. Here, we carried out a standardized comparative study on attention towards a model demonstrating food-related behavior in keas, dogs and children. In a series of experimental sessions, individuals watched different conspecific models while searching, manipulating and feeding. Visual access to the demonstration was provided by two observation holes, which allowed us to determine exactly how often and for how long observers watched the model. We found profound differences in the factors that influence attention within as well as between the tested species. This study suggests that attention should be incorporated as an important variable when testing species in social situations.

Abstract: The question whether a certain species is or is not able to imitate has received much recent attention. However, the ability to copy a demonstrated action might depend not only on the imitative ability of the observer but also on its attention, a factor widely neglected in discussions and experiments. Since attention differs between species as well as between individuals, it is likely to influence the amount and type of information different species and/or observers may extract from a given demonstration. We studied attention in common marmosets, Callithrix jacchus. In a series of experimental sessions, individual marmosets watched different conspecific models that were searching, manipulating an object and feeding. The observers could see the demonstration through two observation holes, which allowed us to measure exactly how often and for how long they watched the model. Marmosets were more attentive towards conspecifics engaged in a problem-solving task than an exploring model. Individual variation in attention was large, ranging from less than 10% to over 49%. Attention also depended on the identity of the model with an increase in attention if it was of the opposite sex. Overall, attention of marmosets was short with a median duration of 6 s while the model manipulated an object. This study measured the real interest of the observer towards a model, which could be an important variable in social-learning experiments.

Abstract: Summary The transmission of cultural knowledge requires learners to identify what relevant information to retain and selectively imitate when observing others' skills. Young human infants--without relying on language or theory of mind--already show evidence of this ability. If, for example, in a communicative context, a model demonstrates a head action instead of a more efficient hand action, infants imitate the head action only if the demonstrator had no good reason to do so, suggesting that their imitation is a selective, interpretative process [1]. Early sensitivity to ostensive-communicative cues and to the efficiency of goal-directed actions is thought to be a crucial prerequisite for such relevance-guided selective imitation [2]. Although this competence is thought to be human specific [2], here we show an analog capacity in the dog. In our experiment, subjects watched a demonstrator dog pulling a rod with the paw instead of the preferred mouth action. In the first group, using the “inefficient” action was justified by the model's carrying of a ball in her mouth, whereas in the second group, no constraints could explain the demonstrator's choice. In the first trial after observation, dogs imitated the nonpreferred action only in the second group. Consequently, dogs, like children, demonstrated inferential selective imitation.