Optimal adult growth of Daphnia in a seasonal environment

1. The cladoceran Daphnia serves as an example of an iteroparous organism, with overlapping generations, that is capable of substantial adult growth. The life history of Daphnia was modelled as the consequence of a series of decisions about allocation of energetic resources to growth and reproduction. 2. We used numerical methods to find resource allocation patterns that maximized fitness of Daphnia in a temporally variable environment. Temporal variation was modelled as alternating active and dormant seasons; length of the active season was uniformly distributed. Fitness was measured by the geometric mean of resting eggs produced at the end of the active season. We examined effects of mean and range of the active season on the optimal life history; we also examined effects of increasing (invertebrate predation), constant (non-selective) and decreasing (fish) size-specific survival rates. For comparison, we found resource allocation patterns that maximized fitness in a constant environment, where fitness was measured by the intrinsic rate of increase r. 3. Life histories optimized for seasonal environments generally showed earlier maturity and greater adult growth than those optimized for constant environments. Adult growth occurred with non-selective predation, and even with fish predation, conditions under which it does not occur in the optimal life histories for constant environments. 4. Greatest size at maturity and adult growth occurred in life histories optimized to invertebrate predation in seasonal environments. Smallest size at maturity and least adult growth occurred in life histories optimized to fish predation. 5. In the optimal life histories, size at maturity generally increased with mean length of the active season. Adult growth reached a maximum for mean seasons of length equal to about one-half to one life span of Daphnia. 6. Increasing the variation in season length decreased adult growth in the optimal life history, but had little effect on size at maturity. 7. We expect that life histories are adapted to the long-term average of season length and its variation. If the animals can detect the type of predator, selection could favour phenotypic variation in resource allocation