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: Animal communication has become a widely studied field of research, especially because of the associated debates on the origin of human language. Due to their phylogenetic proximity with humans, non-human primates represent a suitable model to investigate the precursors of language. This study focuses on the perception of the attentional states of others, an important prerequisite to intentional communication. We investigated whether capuchins (Cebus apella) produce a learnt pointing gesture towards a hidden and unreachable food reward as a function of the attentional status of the human experimenter. For that purpose, we tested five subjects that we first trained to indicate by a pointing gesture towards the human partner the position of a reward hidden by an assistant. Then, capuchins were tested in two experimental conditions randomly ordered. In the first condition—motivation trial—the experimenter was attentive to the subject gestures and rewarded him immediately when it pointed towards the baited cylinder. During the second condition—test trial—the experimenter adopted one of the following attention states and the subject was rewarded after 10 s has elapsed, regardless of the subject’s behaviour. Five attentional states were tested: (1) experimenter absent, (2) experimenter back to the monkey, (3) experimenter’s head away, (4) experimenter watching above the monkey, and (5) experimenter watching the monkey face. Our results reveal a variation in our subjects’ communicative behaviours with a discrimination of the different postural clues (body and head orientation) available in our experimental conditions. This study suggests that capuchins can flexibly use a communicative gesture to adapt to the attentional state of their partner and provides evidence that acquired communicative gestures of monkeys might be used intentionally.

Abstract: Summary For humans, facial expressions are important social signals, and how we perceive specific individuals may be influenced by subtle emotional cues that they have given us in past encounters. A wide range of animal species are also capable of discriminating the emotions of others through facial expressions [1, 2, 3, 4, 5], and it is clear that remembering emotional experiences with specific individuals could have clear benefits for social bonding and aggression avoidance when these individuals are encountered again. Although there is evidence that non-human animals are capable of remembering the identity of individuals who have directly harmed them [6, 7], it is not known whether animals can form lasting memories of specific individuals simply by observing subtle emotional expressions that they exhibit on their faces. Here we conducted controlled experiments in which domestic horses were presented with a photograph of an angry or happy human face and several hours later saw the person who had given the expression in a neutral state. Short-term exposure to the facial expression was enough to generate clear differences in subsequent responses to that individual (but not to a different mismatched person), consistent with the past angry expression having been perceived negatively and the happy expression positively. Both humans were blind to the photograph that the horses had seen. Our results provide clear evidence that some non-human animals can effectively eavesdrop on the emotional state cues that humans reveal on a moment-to-moment basis, using their memory of these to guide future interactions with particular individuals.

Abstract: Summary Domestication has provided the horse with food, shelter, veterinary care and protection, allowing individuals an increased chance of survival. However, the restriction of movement, limited breeding opportunities and a requirement to expend energy, for the benefit of another species, conflict with the evolutionary processes which shaped the behaviour of its predecessors. The behaviour of the horse is defined by its niche as a social prey species but many of the traits which ensured the survival of its ancestors are difficult to accommodate in the domestic environment. There has been a long association between horses and man and many features of equine behaviour suggest a predisposition to interspecific cooperation. However, the importance of dominance in human understanding of social systems has tended to overemphasise its importance in the human-horse relationship. The evolving horse-human relationship from predation to companionship, has resulted in serial conflicts of interest for equine and human participants. Only by understanding the nature and origin of these conflicts can ethologists encourage equine management practices which minimise deleterious effects on the behaviour of the horse.