Abstract: Referential communication occurs when a sender elaborates its gestures to direct the attention of a recipient to its role in pursuit of the desired goal, e.g. by pointing or showing an object, thereby informing the recipient what it wants. If the gesture is successful, the sender and the recipient focus their attention simultaneously on a third entity, the target. Here we investigated the ability of domestic horses (Equus caballus) to communicate referentially with a human observer about the location of a desired target, a bucket of food out of reach. In order to test six operational criteria of referential communication, we manipulated the recipient’s (experimenter) attentional state in four experimental conditions: frontally oriented, backward oriented, walking away from the arena and frontally oriented with other helpers present in the arena. The rate of gaze alternation was higher in the frontally oriented condition than in all the others. The horses appeared to use both indicative (pointing) and non-indicative (nods and shakes) head gestures in the relevant test conditions. Horses also elaborated their communication by switching from a visual to a tactile signal and demonstrated perseverance in their communication. The results of the tests revealed that horses used referential gestures to manipulate the attention of a human recipient so to obtain an unreachable resource. These are the first such findings in an ungulate species.

Abstract: Some domestic animals are thought to be skilled at social communication with humans due to the process of domestication. Horses, being in close relationship with humans, similar to dogs, might be skilled at communication with humans. Previous studies have indicated that they are sensitive to bodily signals and the attentional state of humans; however, there are few studies that investigate communication with humans and responses to the knowledge state of humans. Our first question was whether and how horses send signals to their potentially helpful but ignorant caretakers in a problem-solving situation where a food item was hidden in a bucket that was accessible only to the caretakers. We then examined whether horses alter their behaviours on the basis of the caretakers’ knowledge of where the food was hidden. We found that horses communicated to their caretakers using visual and tactile signals. The signalling behaviour of the horses significantly increased in conditions where the caretakers had not seen the hiding of the food. These results suggest that horses alter their communicative behaviour towards humans in accordance with humans’ knowledge state.

Abstract: Whether non-human animals can recognize human signals, including emotions, has both scientific and applied importance, and is particularly relevant for domesticated species. This study presents the first evidence of horses' abilities to spontaneously discriminate between positive (happy) and negative (angry) human facial expressions in photographs. Our results showed that the angry faces induced responses indicative of a functional understanding of the stimuli: horses displayed a left-gaze bias (a lateralization generally associated with stimuli perceived as negative) and a quicker increase in heart rate (HR) towards these photographs. Such lateralized responses towards human emotion have previously only been documented in dogs, and effects of facial expressions on HR have not been shown in any heterospecific studies. Alongside the insights that these findings provide into interspecific communication, they raise interesting questions about the generality and adaptiveness of emotional expression and perception across species.

Abstract: Quantifying environmental crime and the effectiveness of policy interventions is difficult because perpetrators typically conceal evidence. To prevent illegal uses of natural resources, such as poaching endangered species, governments have advocated granting policy flexibility to local authorities by liberalizing culling or hunting of large carnivores. We present the first quantitative evaluation of the hypothesis that liberalizing culling will reduce poaching and improve population status of an endangered carnivore. We show that allowing wolf (Canis lupus) culling was substantially more likely to increase poaching than reduce it. Replicated, quasi-experimental changes in wolf policies in Wisconsin and Michigan, USA, revealed that a repeated policy signal to allow state culling triggered repeated slowdowns in wolf population growth, irrespective of the policy implementation measured as the number of wolves killed. The most likely explanation for these slowdowns was poaching and alternative explanations found no support. When the government kills a protected species, the perceived value of each individual of that species may decline; so liberalizing wolf culling may have sent a negative message about the value of wolves or acceptability of poaching. Our results suggest that granting management flexibility for endangered species to address illegal behaviour may instead promote such behaviour.

Abstract: Abstract In the study of animal emotions, emotional valence has been found to be difficult to measure. Many studies of farm animals' emotions have therefore focussed on the identification of indicators of strong, mainly negative, emotions. However, subtle variations in emotional valence, such as those caused by rather moderate differences in husbandry conditions, may also affect animals' mood and welfare when such variations occur consistently. In this study, we investigated whether repeated moderate aversive or rewarding events could lead to measurable differences in emotional valence in young, weaned pigs. We conditioned 105 female pigs in a test arena to either a repeated startling procedure (sudden noises or appearances of objects) or a repeated rewarding procedure (applesauce, toy and straw) over 11 sessions. Control pigs were also regularly exposed to the same test arena but without conditioning. Before and after conditioning, we measured heart rate and its variability as well as the behavioural reactions of the subjects in the test arena, with a special focus on detailed acoustic analyses of their vocalisations. The behavioural and heart rate measures were analysed as changes compared to the baseline values before conditioning. A limited number of the putative indicators of emotional valence were affected by the conditioning. We found that the negatively conditioned pigs showed changes that were significantly different from those in control pigs, namely a decrease in locomotion and an increase in standing. The positively conditioned pigs, however, showed a stronger increase in heart rate and a smaller decrease in SDNN (a heart rate variability parameter indicating changes in autonomic regulation) compared to the controls. Compared to the negatively conditioned pigs, the positively conditioned pigs produced fewer vocalisations overall as well as fewer low-frequency grunts but more high-frequency grunts. The low-frequency grunts of the negatively conditioned pigs also showed lower frequency parameters (bandwidth, maximum frequency, 25% and 50% quartiles) compared to those of the positively conditioned pigs. In any of the statistically significant results, the conditioning accounted for 1.5–11.9% of variability in the outcome variable. Hence, we conclude that repeated moderate aversive and rewarding events have weak but measurable effects on some aspects of behaviour and physiology in young pigs, possibly indicating changes in emotional valence, which could ultimately affect their welfare. The combination of ethophysiological indicators, i.e., the concurrent examination of heart rate measures, behavioural responses and especially vocalisation patterns, as used in the current study, might be a useful way of examining subtle effects on emotional valence in further studies.