Abstract: Sixteen domestic dogs (Canis familiaris) were tested in a familiar context in a series of 1-min trials on how well they obeyed after being told by their owner to lie down. Food was used in 1/3 of all trials, and during the trial the owner engaged in 1 of 5 activities. The dogs behaved differently depending on the owner's attention to them. When being watched by the owner, the dogs stayed lying down most often and/or for the longest time compared with when the owner read a book, watched TV, turned his or her back on them, or left the room. These results indicate that the dogs sensed the attentional state of their owners by judging observable behavioral cues such as eye contact and eye, head, and body orientation.

Abstract: The management of the domestic horse often requires them to be kept in isolation from con-specifics. Installing a picture of a horse (generally head and neck view) with a view to providing surrogate companionship has been shown to reduce the negative impact of this isolation. This study aimed firstly to compare the spontaneous response of horses (N=10) to a 2-D image of a horse’s face (FP) with their response to a comparable abstract 2-D image (AP). Secondly, the spontaneous response of horses (N=20) to a 2-D image of a horse’s face with the ears forward (PFP positive) was compared with the response to a 2-D image of a horse’s face with the ears back (NFP negative). The posters were A1 sized and displayed in the horse’s own stable. In study 1, one poster was displayed for 5 minutes and the horse’s behaviour video-recorded. This was removed and the second poster was displayed for 5 minutes and the behaviour video-recorded. FP was displayed first for 5 of the horses and AP displayed first for the other 5. The video footage was observed and the behaviour of the horses and number of times they touched the poster recorded. For the purpose of identifying the area of the poster that was touched by the horse it was divided into 4 equal quarters (TL, TR, BL, BR). In FP the nose of the horse in the 2-D image was located in BL, eyes and ears in TL, chest and lower neck in BR and upper neck in TR. In AP each area contained similar but unique abstract patterns of comparable colour to FP. Differences in behaviour were found according to which poster was displayed. FP was touched significantly more than AP (p=0.001) and was looked at more often (p=0.008). With FP the horses spent significantly longer with their ears forward (p=0.008) and licking and chewing (p=0.016). When the number of touches per poster area was compared (FP and AP) a significant difference was found in the number of times that BL (nose) and BR (chest/lower neck) were touched (p=0.011). Both areas were touched more frequently on FP, with BL being touched the most. In study 2 the same experimental protocol was used to compare responses to positive (PFP) or negative (NFP) 2-D images of a horse’s face (same horse in both PFP and NFP). Again, differences in behaviour were found in response to the two posters. PFP was touched significantly more than NFP (p=0.002) and on both posters the area BL (nose) was touched more frequently than the other areas (PFP: p=0.02, NFP: p=0.01). More ears back behaviour (p<0.001) and more ear locked on behaviour (p=0.008) was shown with NFP. The results of these studies indicate that horses can recognize 2-D images as con-specifics as well as responding to differences in facial expression. There is now the potential for further investigation into the importance of other visual cues in recognition and social interaction as well as the application of findings to enhance equine welfare.

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: The function of olfactory signalling in social species is less well understood than in asocial species. Consequently, we examined olfactory communication in the ringtailed lemur, a socially complex primate that retains a functional vomeronasal organ, has well-developed scent glands and shows a suite of scent-marking behaviour. To assess the information content of different types of scent gland secretions, we decoupled olfactory cues from the visual and behavioural modalities with which scent marking is normally associated. We presented male and female subjects (signal receivers) with a series of choice tests between odours derived from conspecific donors (signal senders) varying by sex, age, social status and reproductive condition. We additionally examined the influence of the receivers' reproductive state and familiarity with the signaller. The reproductive condition, social status and familiarity of senders and receivers affected signal transmission; specifically, male receivers attended most to the odours of conspecifics in breeding condition and to the odours of familiar, dominant animals. By contrast, females varied their responses according to both their own reproductive state and that of the sender. Based on male and female patterns of countermarking, we suggest that scent marking serves a function in intergroup spacing and intrasexual competition for both sexes, as might be expected in a female-dominant species. By contrast, minimal female interest in male odours counters a female mate choice function for scent marking in this species. Nevertheless, scent marks are critical to male-male competition and, therefore, may be subject to sexual selection.

Abstract: Gestures and facial displays are involved in regulating many aspects of mammal social life such as aggression, dominance-subordinate relationships, appeasement and play. Playful activity is an interesting behaviour for examining the role of signals as intentional communication systems. When animals play they perform patterns that are used in other serious contexts. To avoid miscommunication, many species have evolved signals to maintain a playful mood. Bonobos, Pan paniscus, with their flexible social relationships and playful propensity, may represent a good model species to test some hypotheses on adult play signalling. I analysed the potential roles of facial play expressions and solitary play in soliciting and regulating social play and found that adult bonobos used the play face (relaxed open-mouth display) in a selective manner. Play faces were more frequent during social than solitary play and, within social play, polyadic sessions (even though less frequent than dyadic sessions) were characterized by a higher frequency of signals. Following the rule of play intensity matching, play faces were more frequent when the two players matched in age and size (sessions among adults). Moreover, among dyads there was a positive correlation between the frequency of aggressive interactions performed and the frequency of play signals used, thus suggesting that signals are crucial in play negotiations among individuals showing high baseline levels of aggression. Finally, solitary play, especially when it involved pirouettes and somersaults, had an important role in triggering social play.