Abstract
How optimal territory size changes with changes in food density and intruder pressure is in theory complex. Qualitative predictions may change depending upon the goal of the territory holder (time minimization, energy maximization), the possible constraints on energy maximization (time availability, processing capability), and if and how food density and intruder pressure covary. Predictions for the amounts of time spent feeding and defending are similarly variable. Often the qualitative effect of environmental changes will be indeterminate without detailed specification of the functions describing territorial costs and benefits.

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