Christian Nawroth

Farm animal welfare is a major concern for society and food production. To more accurately evaluate animal farming in general and to avoid exposing farm animals to poor welfare situations, it is necessary to understand not only their behavioral but also their cognitive needs and capacities. Thus, general knowledge of how farm animals perceive and interact with their environment is of major importance for a range of stakeholders, from citizens to politicians to cognitive ethologists to philosophers. This review aims to outline the current state of farm animal cognition research and focuses on ungulate livestock species, such as cattle, horses, pigs and small ruminants, and reflects upon a defined set of cognitive capacities (physical cognition: categorization, numerical ability, object permanence, reasoning, tool use; social cognition: individual discrimination and recognition, communication with humans, social learning, attribution of attention, prosociality, fairness). We identify a lack of information on certain aspects of physico-cognitive capacities in most farm animal species, such as numerosity discrimination and object permanence. This leads to further questions on how livestock comprehend their physical environment and understand causal relationships. Increasing our knowledge in this area will facilitate efforts to adjust husbandry systems and enrichment items to meet the needs and preferences of farm animals. Research in the socio-cognitive domain indicates that ungulate livestock possess sophisticated mental capacities, such as the discrimination between, and recognition of, conspecifics as well as human handlers using multiple modalities. Livestock also react to very subtle behavioral cues of conspecifics and humans. These socio-cognitive capacities can impact human-animal interactions during management practices and introduce ethical considerations on how to treat livestock in general. We emphasize the importance of gaining a better understanding of how livestock species interact with their physical and social environments, as this information can improve housing and management conditions and can be used to evaluate the use and treatment of animals during production.

In addition to domestication, interactions with humans or task-specific training during ontogeny have been proposed to play a key role in explaining differences in human-animal communication across species. In livestock, even short-term positive interactions with caretakers or other reference persons can influence human-animal interaction at different levels and over different periods of time. In this study, we investigated human-directed behaviour in the 'unsolvable task' paradigm in two groups of domestic goats (Capra aegagrus hircus). One group was positively handled and habituated to a plastic box by the experimenter to retrieve a food reward, while the other group only received standard husbandry care and was habituated to the box without human assistance. In the unsolvable task, the lid was fixed to the box, with the reward inaccessible to the subjects. The goats were confronted with the unsolvable task three times. We observed no difference between the two groups regarding gaze and contact alternations with the experimenter when confronted with the task they cannot solve by themselves. The goats did not differ in their expression rates of both gaze and contact alternations over three repetitions of the unsolvable task; however, they showed earlier gaze and contact alternations in later trials. The results do not support the hypothesis that short-term positive handling or task-specific training by humans facilitates human-directed behaviour in goats. In contrast, standard husbandry care might be sufficient to establish humans as reference persons for farm animals in challenging situations.

Emotions can be defined as an individual's affective reaction to an external and/or internal event that, in turn, generates a simultaneous cascade of behavioral, physiological, and cognitive changes. Those changes that can be perceived by conspecifics have the potential to also affect other's emotional states, a process labeled as "emotional contagion." Especially in the case of gregarious species, such as livestock, emotional contagion can have an impact on the whole group by, for instance, improving group coordination and strengthening social bonds. We noticed that the current trend of research on emotions in livestock, i.e., investigating affective states as a tool to assess and improve animal welfare, appears to be unbalanced. A majority of studies focuses on the individual rather than the social component of emotions. In this paper, we highlight current limitations in the latter line of research and suggest a stronger emphasis on the mechanisms of how emotions in livestock are transmitted and shared, which could serve as a promising tool to synergistically enhance the welfare of all individuals within a group. Emotions have been defined as short-term affective states elicited by internal and/or external events and are associated with synchronized physiological, behavioral, and cognitive components (1, 2). The framework proposed by Mendl et al. (3) suggests that different aspects of the emotional experience (e.g., neurophysiological, behavioral, and cognitive components) of non-human animals can be assessed along two dimensions, namely valence (negative or positive) and arousal [from low to high (4-6)]. One of the main functions of emotions is to prepare an individual to quickly select an appropriate response (i.e., approach or avoidance depending on the positive or negative valence of the emotion) in order to cope efficiently with its environment (1, 2). Moreover, affective states, i.e., emotions and longer-term affective states (mood), can alter the way in which an individual

Gaze following is widespread among animals. However, the corresponding ultimate functions may vary substantially. Thus, it is important to study previously understudied (or less studied) species to develop a better understanding of the ecological contexts that foster certain cognitive traits. Penguins (Family Spheniscidae), despite their wide interspecies ecological variation, have previously not been considered for cross-species comparisons. Penguin behaviour and communication have been investigated over the last decades, but less is known on how groups are structured, social hierarchies are established, and coordination for hunting and predator avoidance may occur. In this article, we investigated how African penguins (Spheniscus demersus) respond to gaze cues of conspecifics using a naturalistic setup in a zoo environment. Our results provide evidence that members of the family Spheniscidae follow gaze of conspecifics into distant space. However, further tests are necessary to examine if the observed behaviour serves solely one specific function (e.g. predator detection) or is displayed in a broader context (e.g. eavesdropping on relevant stimuli in the environment). In addition, our findings can serve as a starting point for future cross-species comparisons with other members of the penguin family, to further explore the role of aerial predation and social structure on gaze following in social species. Overall, we also suggest that zoo-housed animals represent an ideal opportunity to extend species range and to test phylogenetic families that have not been in the focus of animal cognitive research.