This thesis examines how Eastern grey squirrels, Sciurus carolinensis, modify their foraging and hoarding behaviour in relation to different risks, particularly those which involve a trade-off between securing food resources and avoiding a negative outcome with a competitor. While foraging for food to eat and hoard, squirrels must compete with conspecifics and heterospecifics for access to resources, and they must ensure the safety of their food hoards from onlookers or opportunistic pilferers. While engaging in these behaviours in the most efficient way, they must also avoid being predated upon. Five studies were conducted to further understanding of grey squirrel foraging, hoarding and pilferage behaviours, and how they are affected by different risk factors. The data in this thesis provide experimental evidence that grey squirrels respond directly to conspecific presence as a cue of pilferage risk and adjust their behaviour in ways that may help to reduce cache theft. The data also support the view that conspecific and heterospecific competitors pose risks to foraging and caching, with squirrels modifying their behaviour in ways that serve to avoid negative competitive interactions. Predation risk was found to be particularly disruptive to foraging behaviour, and it also had a seasonal effect upon pilferage rates of experimenter-made caches. A variety of strategies that squirrels might use to pilfer caches were investigated, however, the data did not provide a clear indication of pilferage strategy used by squirrels; they did not seem to use observational spatial memory, and they did not simply pilfer in profitable foraging locations. This thesis raises questions about the mechanisms grey squirrels use to assess pilferage risk and how they engage in pilferage in comparison to other caching species; the studies conducted illustrate different methods that future research could use to investigate food hoarding and pilfering behaviour in wild and captive squirrels.Exeter Graduate Fellowshi
