Energetics, predation, and ration affect size-dependent growth and mortality of fish during winter

Winter temperatures may reduce energy costs for ectotherms. However, variable mid-temperate and low-latitude winters may interact with scaling of size, metabolism, and energy reserves to cause energy deficits and require trade-offs between foraging and predation. A dynamic optimization model explored how ration, fall fat, and both non- and size-selective predation influenced foraging (i.e., fast or forage) and energy allocation (i.e., length or fat) decisions that maximize winter survival of age-0 largemouth bass (Micropterus salmoides). During a mid-latitude (38° N) winter, a pond experiment in which age-0 fish occurred with or without adult conspecific predators tested a subset of the model. In the model without predators, winter foraging occurred, with small size only reducing survival when low ration and low fall fat caused small fish to exhaust reserves. With predation, all sizes foraged to maintain mass and fat reserves when ration was sufficiently high, with small fish also growing in length. When modeled predation was nonselective, size-dependent mortality varied in complex ways. In contrast, size-selective predators consistently reduced survival of small fish. Generally consistent with the model, fish in ponds without predators gained mass and energy content, while those with predators only maintained these parameters. All small individuals grew more than large counterparts in length. Mortality in ponds never depended on size but was ∼20% higher with predators. Energy deficits often demand active foraging during mid-temperate winters, with predation rather than energy depletion influencing size-dependent survival.