Ecological determinants of Cope's rule and its inverse

Cope's rule posits that evolution gradually increases the body size in lineages. Over the last decades, two schools of thought have fueled a debate on the applicability of Cope's rule by reporting empirical evidence, respectively, for and against Cope's rule. The apparent contradictions thus documented highlight the need for a comprehensive process-based synthesis through which both positions of this debate can be understood and reconciled. Here, we use a process-based community-evolution model to investigate the eco-evolutionary emergence of Cope's rule. We report three characteristic macroevolutionary patterns, of which only two are consistent with Cope's rule. First, we find that Cope's rule applies when species interactions solely depend on relative differences in body size and the risk of lineage extinction is low. Second, in environments with higher risk of lineage extinction, the recurrent evolutionary elimination of top predators induces cyclic evolution toward larger body sizes, according to a macroevolutionary pattern we call the recurrent Cope's rule. Third, when interactions between species are determined not only by their body sizes but also by their ecological niches, the recurrent Cope's rule may get inverted, leading to cyclic evolution toward smaller body sizes. This recurrent inverse Cope's rule is characterized by highly dynamic community evolution, involving the diversification of species with large body sizes and the extinction of species with small body sizes. To our knowledge, these results provide the first theoretical foundation for reconciling the contrasting empirical evidence reported on body-size evolution.