The temperature-size rule in ectotherms: simple evolutionary explanations may not be general

Abstract

In many organisms, individuals in colder environments grow more slowly but are larger as adults. This widespread pattern is embodied by two well-established rules: Bergmann's rule, which describes the association between temperature and body size in natural environments, and the temperature-size rule, which describes reaction norms relating temperature to body size in laboratory experiments. Theory predicts that organisms should grow to be larger in colder environments when growth efficiency decreases with increasing environmental temperature. Using data from 97 laboratory experiments, including 58 species of ectotherms, we found little evidence that growth efficiency is negatively related to environmental temperature within the thermal range that is relevant to the temperature-size rule. Instead, growth efficiency was either positively related or insensitive to environmental temperature in the majority of cases (73 of 89 cases for gross growth efficiency and 18 of 24 cases for net growth efficiency). Two possibilities merit consideration. First, high temperatures may impose constraints on growth that only arise late during ontogeny; this simple and potentially general explanation is supported by the fact that thermal optima for growth efficiency and growth rate decrease as individuals grow. Alternatively, the general explanation for relationships between temperature and body size may not be simple. If the latter view is correct, the best approach might be to generate and test theories that are tailored specifically to organisms with similar behavior and physiology.