Document Type


Date of Original Version



Biological Sciences


Many crustaceans have complex life cycles characterized by a metamorphosis, yet variation in metamorphic traits, and the causes and consequences of this variation, have rarely been examined. Food concentrations were changed during specific larval stages of the freshwater copepod Mesocyclops edax Forbes (Copepoda: Cyclopoida) to examine whether age and size at metamorphosis remain flexible or become fixed during the larval period. Results were compared to predictions of both flexible (the Wilbur—Collins model) and fixed (Leips—Travis model) rate models for the timing of amphibian metamorphosis. Age and size at metamorphosis were variable in all treatments, and age was always more variable than size. Changes in food concentration early in larval development resulted in significant differences in age at metamorphosis among treatments, but changes initiated when 60% of the larval period had passed had no effect on age at metamorphosis. Development appeared to become fixed later in the larval period, before the ultimate larval stage was reached. These results support predictions of the Leips—Travis model. Early changes in food concentrations had significant effects on size at metamorphosis, but changes initiated during the penultimate larval stage (50—60% of larval development) had no effect on metamorph size. Size at metamorphosis in M. edax also appeared to be fixed before the ultimate larval stage was reached. Fixation of size at metamorphosis during development is not predicted by either model and may be unique to organisms with rigid exoskeletons that constrain growth within any stage. Patterns of covariation between age and size at metamorphosis suggest that food conditions early in larval development exert a large effect on metamorphic traits, in contrast to patterns observed in several amphibian species. The Wilbur—Collins model places a fitness premium on delaying metamorphosis to achieve a maximum size, when growth conditions are favorable; it thus may not apply to crustaceans. Selection pressures on the timing of metamorphosis in crustaceans may differ substantially from those identified for amphibians and other organisms. Because of these differences, incorporating crustaceans into studies of metamorphosis will help to clarify the factors affecting this life cycle transition.