Dither: A unifying model of the effects of visitor numbers on zoo animal behavior.

Interest in the impact of human presence on the behavior and well-being of zoo and aquarium animals is increasing. Previous work has conceptualized the presence of zoo visitors as having one of three impacts on the behavior of animals in zoos: positive, negative, or neutral. Research suggests the same species may exhibit all three responses under different conditions, calling into question whether the positive/negative/neutral framework is the most useful way of considering visitor impact on animal behavior. Here we present a model of visitor effects that unifies these three predictions. Our model suggests that zoo-goers may provide a "dither effect" for some animals living in zoos. We posit animals may show nonlinear behavioral responses over a range of visitor densities, effectively exhibiting changes in both comfortable and anxiety-like behaviors under different levels of human presence. We tested this model during two COVID-19 related closures at the San Francisco Zoo, studying seven species for evidence of nonlinear relationships between visitor numbers and animal behavior. Our results support the dither effect acting in several species observed.

Space, time, and context drive anticipatory behavior: Considerations for understanding the behavior of animals in human care.

Animal-based measures reflecting the welfare state of individuals are critical for ensuring the well-being of animals under human care. Anticipatory behavior is one potential animal-based measure that has gained traction in recent years, as it is theorized to relate to animals' reward sensitivity. It is of particular interest as an assessment for animals living under human care, as the predictability of the captive environment lends itself to the development of this class of behaviors. Animals are likely to exhibit anticipation in locations related to the anticipated event, often in temporally predictable time frames, and before specific contexts they experience in their day-to-day management. In this sense and under certain circumstances, anticipatory behaviors are likely to drive observed behavioral or space use patterns of animals under human care. Drawing conclusions from such data without identifying anticipation may result in misleading conclusions. Here we discuss how space, time, and context are related to patterns of anticipatory behaviors in animals under human care, how unidentified anticipation may alter conclusions regarding animal behavior or welfare under certain circumstances.