Abstract

In the wild, primates are selective over the routes that they take when foraging and seek out preferred or ephemeral food. Given this, we tested how a group of captive chimpanzees weighed the relative benefits and costs of foraging for food in their environment when a less-preferred food could be obtained with less effort than a more-preferred food. In this study, a social group of six zoo-housed chimpanzees (Pan troglodytes) could collect PVC tokens and exchange them with researchers for food rewards at one of two locations. Food preference tests had revealed that, for these chimpanzees, grapes were a highly-preferred food while carrot pieces were a less-preferred food. The chimpanzees were tested in three phases, each comprised of 30 thirty-minute sessions. In phases 1 and 3, if the chimpanzees exchanged a token at the location they collected them they received a carrot piece (no travel) or they could travel ≥10 m to exchange tokens for grapes at a second location. In phase 2, the chimpanzees had to travel for both rewards (≥10 m for carrot pieces, ≥15 m for grapes). The chimpanzees learned how to exchange tokens for food rewards, but there was individual variation in the time it took for them to make their first exchange and to discover the different exchange locations. Once all the chimpanzees were proficient at exchanging tokens, they exchanged more tokens for grapes (phase 3). However, when travel was required for both rewards (phase 2), the chimpanzees were less likely to work for either reward. Aside from the alpha male, all chimpanzees exchanged tokens for both reward types, demonstrating their ability to explore the available options. Contrary to our predictions, low-ranked individuals made more exchanges than high-ranked individuals, most likely because, in this protocol, chimpanzees could not monopolize the tokens or access to exchange locations. Although the chimpanzees showed a preference for exchanging tokens for their more-preferred food, they appeared to develop strategies to reduce the cost associated with obtaining the grapes, including scrounging rewards and tokens from group mates and carrying more than one token when travelling to the farther exchange location. By testing the chimpanzees in their social group we were able to tease apart the social and individual influences on their decision making and the interplay with the physical demands of the task, which revealed that the chimpanzees were willing to travel farther for better.

Highlights

  • How, when, where, and for how long animals forage for food is influenced by external factors, internal factors, and phylogenetic factors (Pianka, 1997)

  • Observations of wild black howler monkeys (Alouatta pigra) suggests that their foraging strategy is influenced by both distance and food desirability (they use step-wise movements to reach high quality patches, (Plante, Colchero & Calme, 2014))

  • There was no correlation between the chimpanzees’ rank and the order in which they acquired this task; there was a negative correlation between rank and number of tokens exchanged across all sessions (Spearmans’s rho: rs = −1.00, N = 6, P < 0.001)

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Summary

Introduction

How, when, where, and for how long animals forage for food is influenced by external factors (e.g., predation risk, social interactions, and prey availability), internal factors (e.g., hunger, food preferences, and animals’ age or sex), and phylogenetic factors (e.g., physiological parameters and sensory limitations) (Pianka, 1997). In order to obtain preferred or ephemeral foods, primates are selective about which trees they feed from (Glander , 1979) and the routes that they travel to reach them (Ban, Boesch & Janmaat, 2014). Observations of wild black howler monkeys (Alouatta pigra) suggests that their foraging strategy is influenced by both distance (they preferentially travel to closer trees) and food desirability (they use step-wise movements to reach high quality patches, (Plante, Colchero & Calme, 2014)). Female chimpanzees (Pan troglodytes) have been reported to leave their nighttime nests earlier when more-preferred fig trees are located farther away (Janmaat et al, 2014), indicating that they are willing to travel longer distances to reach more desirable food rewards and that they plan their activities in order to reach them. Even in a captive setting, where primates are not required to forage for their daily food to survive, they have been reported to use efficient routes when searching for foods, sometimes bypassing less-preferred foods to reach more-preferred foods first (e.g., Menzel, 1973; Boesch & Boesch, 1984 provides a review), but evidence for such strategic foraging is mixed (e.g., Howard & Fragaszy, 2014)

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