Abstract

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.

Highlights

  • Humans have had a negative impact on other animals for millennia (Barnosky et al, 2004) and, with the human population continuing to grow (Roser et al, 2013), wild animals may encounter humans with increasing frequency

  • As humans are a key driver of wildlife declines, understanding the behavioral and cognitive processes that shape wild animals’ responses to humans is likely to be important in mitigating the detrimental effects of human activity

  • To successfully navigate encounters with humans, animals rely on a wide range of cognitive processes, as they must perceive and attend to relevant cues, integrate this information with previous experience, and mount the appropriate behavioral response (Figure 1)

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Summary

INTRODUCTION

Humans have had a negative impact on other animals for millennia (Barnosky et al, 2004) and, with the human population continuing to grow (Roser et al, 2013), wild animals may encounter humans with increasing frequency. As domesticated animals have been selected for docility and sociability toward humans (Wilkins et al, 2014; VonHoldt et al, 2017), the responses of such animals, even when feral, are likely to differ substantially from those of species with no evolutionary history of domestication. Perhaps the best known case of humanmediated extinction in modern history is that of the dodo (Raphus cucullatus), whose naïveté to predators rendered the species vulnerable to exploitation by humans (Cheke, 2004). To this day, wild animals risk following the same fate (Ripple et al, 2019). We conclude by emphasizing the important role of animal cognition research in reducing human-wildlife conflict and improving conservation outcomes

HOW DO WILD ANIMALS RESPOND TO HUMANS AS A SPECIES?
Do Wild Animals Perceive Humans as Causal Agents?
Dangerous Humans
Neutral Humans
Rewarding Humans
HOW DO WILD ANIMALS DISTINGUISH BETWEEN DANGEROUS AND NEUTRAL HUMANS?
Gaze Direction
INDIVIDUAL RECOGNITION OF HUMANS
Social Learning About Dangerous Individual Humans
VARIATION IN RESPONSES TO HUMANS
Why Do Animals Vary in Their Responses to Humans?
Wider Implications
CONSERVATION AND MANAGEMENT APPLICATIONS
FUTURE DIRECTIONS
AUTHOR CONTRIBUTIONS
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