Mechanistic Temperature-Size Rule Explanation Should Reconcile Physiological and Mortality Responses to Temperature

AbstractThe temperature-size rule is one of the universal rules in ecology and states that ectotherms in warmer waters will grow faster as juveniles, mature at smaller sizes and younger ages, and reach smaller maximum body sizes. Many models have unsuccessfully attempted to reproduce temperature-size rule-consistent life histories by using two-term (anabolism and catabolism) Pütter-type growth models, such as the von Bertalanffy. Here, we present a physiologically structured individual growth model, which incorporates an energy budget and optimizes energy allocation to growth, reproduction, and reserves. Growth, maturation, and reproductive output emerge as a result of life-history optimization to specific physiological rates and mortality conditions. To assess which processes can lead to temperature-size rule-type life histories, we simulate 42 scenarios that differ in temperature and body size dependencies of intake, metabolism, and mortality rates. Results show that the temperature-size rule can emerge in two ways. The first way requires both intake and metabolism to increase with temperature, but the temperature-body size interaction of the two rates must lead to relatively faster intake increase in small individuals and relatively larger metabolism increase in large ones. The second way requires only higher temperature-driven natural mortality and faster intake rates in early life (no change in metabolic rates is needed). This selects for faster life histories with earlier maturation and increased reproductive output. Our model provides a novel mechanistic and evolutionary framework for identifying the conditions necessary for the temperature-size rule. It shows that the temperature-size rule is likely to reflect both physiological changes and life-history optimization and that use of von Bertalanffy-type models, which do not include reproduction processes, can hinder our ability to understand and predict ectotherm responses to climate change.