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Humans have a profound effect on the planet’s ecosystems, and unprecedented rates of human population growth and urbanization have brought wild animals into increasing contact with people. For many species, appropriate responses toward humans are likely to be critical to survival and reproductive success. Although numerous studies have investigated the impacts of human activity on biodiversity and species distributions, relatively few have examined the effects of humans on the behavioral responses of animals during human-wildlife encounters, and the cognitive processes underpinning those responses. Furthermore, while humans often present a significant threat to animals, the presence or behavior of people may be also associated with benefits, such as food rewards. In scenarios where humans vary in their behavior, wild animals would be expected to benefit from the ability to discriminate between dangerous, neutral and rewarding people. Additionally, individual differences in cognitive and behavioral phenotypes and past experiences with humans may affect animals’ ability to exploit human-dominated environments and respond appropriately to human cues. In this review, we examine the cues that wild animals use to modulate their behavioral responses toward humans, such as human facial features and gaze direction. We discuss when wild animals are expected to attend to certain cues, how information is used, and the cognitive mechanisms involved. We consider how the cognitive abilities of wild animals are likely to be under selection by humans and therefore influence population and community composition. We conclude by highlighting the need for long-term studies on free-living, wild animals to fully understand the causes and ecological consequences of variation in responses to human cues. The effects of humans on wildlife behavior are likely to be substantial, and a detailed understanding of these effects is key to implementing effective conservation strategies and managing human-wildlife conflict.

Humans have a profound effect on the planet's ecosystems, and unprecedented rates of human population growth and urbanization have brought wild animals into increasing contact with people. For many species, appropriate responses toward humans are likely to be critical to survival and reproductive success. Although numerous studies have investigated the impacts of human activity on biodiversity and species distributions, relatively few have examined the effects of humans on the behavioral responses of animals during human-wildlife encounters, and the cognitive processes underpinning those responses. Furthermore, while humans often present a significant threat to animals, the presence or behavior of people may be also associated with benefits, such as food rewards. In scenarios where humans vary in their behavior, wild animals would be expected to benefit from the ability to discriminate between dangerous, neutral and rewarding people. Additionally, individual differences in cognitive and behavioral phenotypes and past experiences with humans may affect animals' ability to exploit human-dominated environments and respond appropriately to human cues. In this review, we examine the cues that wild animals use to modulate their behavioral responses toward humans, such as human facial features and gaze direction. We discuss when wild animals are expected to attend to certain cues, how information is used, and the cognitive mechanisms involved. We consider how the cognitive abilities of wild animals are likely to be under selection by humans and therefore influence population and community composition. We conclude by highlighting the need for long-term studies on free-living, wild animals to fully understand the causes and ecological consequences of variation in responses to human cues. The effects of humans on wildlife behavior are likely to be substantial, and a detailed understanding of these effects is key to implementing effective conservation strategies and managing human-wildlife conflict. [ABSTRACT FROM AUTHOR]

Gaze following is the ability to utilise information from another's gaze. It is most often seen in a social context or as a reflexive response to interesting external stimuli. Social species can potentially reveal utilisable knowledge about another's future intentions by attending to the target of their gaze. However, in even more fundamental situations, being sensitive to another's gaze can also be useful such as when it can facilitate greater foraging efficiency or lead to earlier predator detection. While gaze sensitivity has been shown to be prevalent in a number of social species, little is currently known about the potential for gaze following in asocial species. The current study investigated whether an asocial reptile, the leopard gecko (Eublepharis macularius), could reliably use the visual indicators of attention to follow the gaze of a conspecific around a barrier. We operated three trial conditions and found subjects (N = 6) responded significantly more to the conspecific demonstrator looking up at a laser stimulus projected onto an occluder during the experimental condition compared to either of two control conditions. The study's findings point toward growing evidence for gaze-following ability in reptiles, who are typically categorised as asocial. Furthermore, our findings support developing comparative social cognition research showing the origins of gaze following and other cognitive behaviours that may be more widely distributed across taxonomic groups than hitherto thought. [ABSTRACT FROM AUTHOR]

Attending to where others are looking is thought to be of great adaptive benefit for animals when avoiding predators and interacting with group members. Many animals have been reported to respond to the gaze of others, by co-orienting their gaze with group members (gaze following) and/or responding fearfully to the gaze of predators or competitors (i.e., gaze aversion). Much of the literature has focused on the cognitive underpinnings of gaze sensitivity, namely whether animals have an understanding of the attention and visual perspectives in others. Yet there remain several unanswered questions regarding how animals learn to follow or avoid gaze and how experience may influence their behavioral responses. Many studies on the ontogeny of gaze sensitivity have shed light on how and when gaze abilities emerge and change across development, indicating the necessity to explore gaze sensitivity when animals are exposed to additional information from their environment as adults. Gaze aversion may be dependent upon experience and proximity to different predator types, other cues of predation risk, and the salience of gaze cues. Gaze following in the context of information transfer within social groups may also be dependent upon experience with group-members; therefore we propose novel means to explore the degree to which animals respond to gaze in a flexible manner, namely by inhibiting or enhancing gaze following responses. We hope this review will stimulate gaze sensitivity research to expand beyond the narrow scope of investigating underlying cognitive mechanisms, and to explore how gaze cues may function to communicate information other than attention. [ABSTRACT FROM AUTHOR]

Predator recognition is a prerequisite for antipredator behaviour. Although species level predator recognition is well documented, there is emerging evidence that some birds, including corvids, can differentiate between individual humans that pose different levels of threat. Other predator cues such as gaze direction may offer additional important information regarding the likelihood of an attack, but it is unknown whether wild animals can integrate information about the individual identity and gaze direction of a predator when perceiving threat levels. Here we measured wild jackdaws' responses to human head orientation, directed towards or away from their nestbox containing chicks. To test whether aversive responses to gaze direction were integrated with information about human identity, the human presenting gaze cues wore one of two masks: a ‘threatening’ mask that had previously been worn when handling the jackdaws' chicks and a ‘neutral mask’ that had previously been worn when walking by, but not disturbing the nest. Latency to return to the nestbox was significantly higher when the threatening mask rather than the neutral mask was worn, but it was not influenced by head direction. However, once a jackdaw landed outside the nestbox, there was a nonsignificant trend for it to be quicker to enter the box when the human was gazing towards, rather than away from the box. These results indicate that wild jackdaws can recognize different individual humans and thus differentiate between those wearing a threatening and a neutral mask, and respond defensively by guarding their chicks from potential threat. Jackdaws may integrate both predator identity and gaze cues; however, predation risk is primarily perceived through individual human recognition. [ABSTRACT FROM AUTHOR]

Sensitivity to the gaze of other individuals has long been a primary focus in sociocognitive research on humans and other animals. Information about where others are looking may often be of adaptive value in social interactions and predator avoidance, but studies across a range of taxa indicate there are substantial differences in the extent to which animals obtain and use information about other individuals' gaze direction. As the literature expands, it is becoming increasingly difficult to make comparisons across taxa as experiments adopt and adjust different methodologies to account for differences between species in their socioecology, sensory systems and possibly also their underlying cognitive mechanisms. Furthermore, as more species are found to exhibit gaze sensitivity, more terminology arises to describe the behaviours. To clarify the field, we propose a restricted nomenclature that defines gaze sensitivity in terms of observable behaviour, independent of the underlying mechanisms. This is particularly useful in nonhuman animal studies where cognitive interpretations are ambiguous. We then describe how socioecological factors may influence whether species will attend to gaze cues, and suggest links between ultimate factors and proximate mechanisms such as cognition and perception. In particular, we argue that variation in sensory systems, such as retinal specializations and the position of the eyes, will determine whether gaze cues (e.g. head movement) are perceivable during visual fixation. We end by making methodological recommendations on how to apply these variations in socioecology and visual systems to advance the field of gaze research. [Copyright &y& Elsevier]

Abstract: Eye contact is important to share communication during social interactions. However, how accurately humans can perceive the gaze direction of others toward themselves and whether pupils dilate when humans consciously or unconsciously perceive own eyes are looked by others remain unclear. In this study, we examined the relationship between the explicit perception of looking into each other''s eyes and the implicit physiological response of pupillary dilation by using an original face-to-face method. We found that humans do not correctly detect the gaze direction of others. Furthermore, one''s pupils dilated when one gazed at others’ eyes. Awareness of others’ gaze on one''s eyes, rather than the actual focusing of other''s gaze on one''s eyes, enhanced pupillary dilation. Therefore, physiological responses are caused not when people actually look into each other''s gaze, but when the consciousness of other''s gaze is activated, which suggests that eye contact often involves one-way communication. [Copyright &y& Elsevier]

Human-wildlife conflict is one of the greatest threats to species populations worldwide. One species facing national declines in the UK is the herring gull (Larus argentatus), despite an increase in numbers in urban areas. Gulls in urban areas are often considered a nuisance owing to behaviours such as food-snatching. Whether urban gull feeding behaviour is influenced by human behavioural cues, such as gaze direction, remains unknown. We therefore measured the approach times of herring gulls to a food source placed in close proximity to an experimenter who either looked directly at the gull or looked away. We found that only 26% of targeted gulls would touch the food, suggesting that food-snatching is likely to be conducted by a minority of individuals. When gulls did touch the food, they took significantly longer to approach when the experimenter's gaze was directed towards them compared with directed away. However, inter-individual behaviour varied greatly, with some gulls approaching similarly quickly in both treatments, while others approached much more slowly when the experimenter was looking at them. These results indicate that reducing human-herring gull conflict may be possible through small changes in human behaviour, but will require consideration of behavioural differences between individual gulls.

Looking where others are allocating attention can facilitate social interactions by providing information about objects or locations of interest. We asked whether European starlings follow the orientation behaviour of conspecifics owing to their highly gregarious behaviour. Starlings reoriented their attention to follow that of a robot around a barrier more often than when the robot's attention was directed elsewhere. This is the first empirical evidence of reorienting in response to conspecific attention in a songbird. Starlings may use this behaviour to obtain fine-tuned spatial information from conspecifics (e.g. direction of predator approach, spatial location of food patches), enhancing group cohesion., (© 2014 The Author(s) Published by the Royal Society. All rights reserved.)