Abstract: Abstract  Great ape gestures have attracted considerable research interest in recent years, prompted by their flexible and intentional pattern of use; but almost all studies have focused on single gestures. Here, we report the first quantitative analysis of sequential gesture use in western gorillas (Gorilla gorilla gorilla), using data from three captive groups and one African study site. We found no evidence that gesture sequences were given for reasons of increased communicative efficiency over single gestures. Longer sequences of repeated gestures did not increase the likelihood of response, and using a sequence was seldom in reaction to communicative failure. Sequential combination of two gestures with similar meanings did not generally increase effectiveness, and sometimes reduced it. Gesture sequences were closely associated with play contexts. Markov transition analysis showed two networks of frequently co-occurring gestures, both consisting of gestures used to regulate play. One network comprised only tactile gestures, the other a mix of silent, audible and tactile gestures; apparently, these clusters resulted from gesture use in play with proximal or distal contact, respectively. No evidence was found for syntactic effects of sequential combination: meanings changed little or not at all. Semantically, many gestures overlapped massively with others in their core information (i.e. message), and gesture messages spanned relatively few functions. We suggest that the underlying semantics of gorilla gestures is highly simplified compared to that of human words. Gesture sequences allow continual adjustment of the tempo and nature of social interactions, rather than generally conveying semantically referential information or syntactic structures.

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: Equine vocalization and acoustic sounds can communicate a horse’s emotional state, physiological state, and situation to other individuals, including other horses and humans. These vocalizations and acoustic sounds can be divided into several types. The whinny, nicker, squeal, blow, snore, snort, roar, and groan are typical types of horse vocalizations and acoustic sounds. The sound localization thresholds of horses are markedly poorer than those of other large mammals, such as humans and elephants. The audiogram of horse has shown their best sensitivity and hearing range in which it perceives sound. Laryngeal diseases, such as laryngeal hemiplegia, dorsal displacement of the soft palate, and alar fold paralysis, can cause laryngeal sounds in the upper airway. The analyses of horses’ vocalizations and laryngeal sounds that are reviewed in this article were conducted with computer-aided analysis programs using spectrograms and spectra that evaluate several parameters, including amplitude, fundamental frequency, duration, and formants. Laryngeal sound analysis could be a useful method for diagnosing upper airway diseases. This article presents a review of the literature describing scientific analyses of horse vocalizations and acoustic sounds to elucidate equine acoustic communications and aid in the development of horse-human bonds.

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: 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.