Modeling Global Macroclimatic Constraints on Ectotherm Energy Budgets

We describe a mechanistic individual-based model of how globalmacroclimatic constraints affect the energy budgets of ectothermicanimals. The model uses macroclimatic and biophysical charactersof the habitat and organism and tenets of heat transfer theoryto calculate hourly temperature availabilities over a year.Data on the temperature dependence of activity rate, metabolism,food consumption and food processing capacity are used to estimatethe net rate of resource assimilation which is then integratedover time. We detail and explore the significance of assumptionsused in these calculations. We present a new test of this model in which we show that thepredicted energy budget sizes for 11 populations of the lizardSceloporus undulalus are in close agreement with observed resultsfrom previous field studies. This demonstrates that model testsare feasible and the results are reasonable. Further, sincethe model represents an upper bound to the size of the energybudget, observed residual deviations form explicit predictionsabout the effects of environmental constraints on the bioenergeticsof the study lizards within each site that may be tested byfuture field and laboratory studies. Three major new improvements to our modeling are discussed.We present a means to estimate microclimate thermal heterogeneitymore realistically and include its effects on field rates ofindividual activity and food consumption. Second, we describean improved model of digestive function involving batch processingof consumed food. Third, we show how optimality methods (specificallythe methods of stochastic dynamic programming) may be includedto model the fitness consequences of energy allocation decisionssubject to food consumption and processing constraints whichare predicted from the microclimate and physiological modeling Individual-based models that incorporate macroclimatic constraintson individual resource acquisition, assimilation and allocationcan provide insights into theoretical investigations about theevolution of life histories in variable environments as wellas provide explicit predictions about individual, populationand community level responses to global climate change.