Complex Life Cycles in a Variable Environment: Predicting When the Timing of Metamorphosis Shifts from Resource Dependent to Developmentally Fixed.

To test models of the timing of and size at metamorphosis, researchers manipulate food at several times during the larval phase of an animal's complex life cycle. Data from diverse taxa show that the age at metamorphosis becomes resource independent (i.e., fixed) at some point during the larval phase. Although existing models have been modified to incorporate a fixed rate of development, none predicts when phenotypic plasticity in metamorphic timing is lost. A graphical model is presented that extends knowledge of a genotype's optimal age and size at metamorphosis in different environments in which resources remain constant throughout the larval phase (i.e., the genotype's reaction norm) to predict when development rate becomes fixed in response to resource variability during the larval phase. Model predictions concur with data from food-switching experiments on anuran tadpoles and barnacle nauplii. As interest in the timing of and size at metamorphosis expands from well-studied taxa (e.g., amphibians) to the many others that have complex life cycles, the predictive model provides a useful tool to design and improve experiments.