How do female desert tortoises (Gopherus agassizii) reproduce every year despite variability in winter rainfall and food availability? To answer this question, I measured energy budgets of individual female desert tortoises from July 1987 to July 1989. Females produced eggs in years with low levels of winter annual plants by relaxing their control of energy and water homeostasis. They tolerated large deficits and surpluses in their body dry-matter composition (both nonlipid and lipid) on a seasonal, annual, and longer time scale. They could increase body energy content (lipid and nonlipid energy) before winter and use this reserve (especially nonlipid energy) the following spring to produce eggs. Females used high-protein foods and rainwater, when available, to achieve energy surpluses that helped them survive periods of low resource availability (e.g., during hibernation and droughts). Adjusting seasonal and annual field metabolic rates (FMR) and food requirements to levels of food availability, they sti...