Seasonal changes in territory use by red squirrels, Tamiasciurus hudsonicus, and responses to food augmentation
Traditionally, the importance of food availability and intruder pressure on habitat use have been investigated through their influences on territory size. Food-augmentation studies are more prevalent; however, they are potentially confounded by the indirect effect of conspecific attraction to local food additions. Frequent ambiguous results may be attributed to the scale of investigation. Furthermore, such investigations have not considered potential seasonal influences. In this study, controlled experimental manipulations of food availability and conspecific density were performed within red squirrel (Tamiasciurus hudsonicus) territories in Algonquin Park, Ontario, to tease apart the relative importance of direct and indirect effects of food augmentation. Effects were investigated at two levels of scale, using territory size and giving-up densities (GUDs) within activity locations as response variables. Augmenting food, with or without simultaneously removing conspecifics, did not influence territory size. However, territory size decreased in control and treatment groups following manipulations, which is suggestive of seasonal influences associated with caching behaviour and midden defence. Despite overwhelming seasonal influences, GUDs were more sensitive to treatments, revealing that competitors had a greater influence on habitat use than the direct effect of food addition. Ambiguous results of previous studies may due to the use of territory-size estimators that are insensitive to responses occurring at smaller scales.