In territorial species, competitive asymmetries can result in the uneven distribution of food resources as high-quality individuals force young or subordinates to occupy smaller or lower-quality sites, or both. However, spatiotemporal variation in the production of resources also can influence an individual's ability to monopolize resources and, consequently, affect survival. We examined how spatial and temporal variation in food supply affects the distribution of resources among territorial food-hoarding red squirrels (Tamiasciurus hudsonicus) by mapping cone production over 62.5 ha of white spruce (Picea glauca) forest and measuring the interrelationships between territory size, cone supply, and survival during 4 years of low cone abundance in Yukon, Canada. Territory size and cone production within a territory varied 10- and 520-fold, respectively, with juvenile squirrels occupying smaller territories with fewer cones. Because of small-scale heterogeneity in the distribution of cones, territory size explained low to moderate amounts of the variation in territory-wide cone production within (rs = 0.49–0.69) and across (rs = 0.31) years. Furthermore, spatial heterogeneity in cone production varied across years such that territory location was not useful for predicting relative food availability from one year to the next. Perhaps as a consequence of this heterogeneity, the number of cones within an individual's hoard was only weakly correlated with territory size (rs = 0.42) and not correlated with territory-wide cone production. Hoard size better predicted overwinter survival than did territory size, whereas cone production on a territory was not predictive of survival. We suggest that caution be used in equating territory size with food supply because small-scale spatial heterogeneity can weaken this relationship, particularly in food-hoarders. Spatiotemporal variation in food abundance, in turn, may cause high-quality territories in one year to be low-quality territories in other years, which will limit the ability of individuals to assess the long-term quality of territories at the time of settlement.
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Vol. 94 • No. 5