Primate cognition research is reliant on access to members of the study sp ecies and logistical infrastructures to conduct observations and experiments. Historically founded in research centers and private collections, and spreading to modern zoos, sanctuaries, and the field, primate cognition has been investigated in diverse settings, each with benefits and challenges. In our systematic review of 12 primatology, animal behavior, and animal cognition journals over the last 15 years, we turn a spotlight on zoos to quantify their current impact on the field and to highlight their potential as robust contributors to future work. To put zoo-based research in context, we compare zoos to three other site types: university-owned or independent research centers, sanctuaries, and field sites. We assess the contributions of zoos across several critical considerations in primate cognition research, including number of investigations, species diversity, sample size, research topic diversity, and methodology. We identified 1119 publications reporting studies of primate cognition, almost 25% of which report research conducted in zoos. Across publications, zoobased research has greater species diversity than research centers and covers a diverse range of research topics. Although our review is merely a snapshot of primate cognition research, our findings suggest that zoos may present advantages to researchers regarding species diversity, and lack some of the methodological constraints of field sites, allowing greater ease of access to a diverse range of subjects for cognition investigations. We suggest that zoos have great potential as key contributors for future investigations in primate cognition. Finally, we shed light on the symbiotic relationship that can emerge between researchers and zoos, forming partnerships that bring unique advantages to both parties.

Almost all animals navigate their environment to find food, shelter, and mates. Spatial cognition of nonhuman primates in large-scale environments is notoriously difficult to study. Field research is ecologically valid, but controlling confounding variables can be difficult. Captive research enables experimental control, but space restrictions can limit generalizability. Virtual reality technology combines the best of both worlds by creating large-scale, controllable environments. We presented six chimpanzees with a seminaturalistic virtual environment, using a custom touch screen application. The chimpanzees exhibited signature behaviors reminiscent of real-life navigation: They learned to approach a landmark associated with the presence of fruit, improving efficiency over time; they located this landmark from novel starting locations and approached a different landmark when necessary. We conclude that virtual environments can allow for standardized testing with higher ecological validity than traditional tests in captivity and harbor great potential to contribute to longstanding questions in primate navigation, e.g., the use of landmarks, Euclidean maps, or spatial frames of reference.

Humans can tell when they find a task difficult. Subtle uncertainty behaviors like changes in motor speed and muscle tension precede and affect these experiences. Theories of animal metacognition likewise stress the importance of endogenous signals of uncertainty as cues that motivate metacognitive behaviors. However, while researchers have investigated second-order behaviors like information seeking and declining difficult trials in nonhuman animals, they have devoted little attention to the behaviors that express the cognitive conflict that gives rise to such behaviors in the first place. Here we explored whether three chimpanzees would, like humans, show hand wavering more when faced with more difficult choices in a touch screen transitive inference task. While accuracy was very high across all conditions, all chimpanzees wavered more frequently in trials that were objectively more difficult, demonstrating a signature behavior which accompanies experiences of difficulty in humans. This lends plausibility to the idea that feelings of uncertainty, like other emotions, can be studied in nonhuman animals. We propose to routinely assess uncertainty behaviors to inform models of procedural metacognition in nonhuman animals.