Tempo and mode in evolution: phylogenetic inertia, adaptation and comparative methods

Before the Evolutionary Synthesis, 'phylogenetic inertia' was associated with theories of orthogenesis, which claimed that organisms possessed an endogenous perfecting principle. The concept in the modern literature dates to Simpson (1944), who used 'evolutionary inertia' as a description of pattern in the fossil record. Wilson (1975) used 'phylogenetic inertia' to describe population-level or organismal properties that can affect the course of evolution in response to selection. Many current authors now view phylogenetic inertia as an alternative hypothesis to adaptation by natural selection when attempting to explain interspecific variation, covariation or lack thereof in phenotypic traits. Some phylogenetic comparative methods have been claimed to allow quantification and testing of phylogenetic inertia. Although some existing methods do allow valid tests of whether related species tend to resemble each other, which we term 'phylogenetic signal', this is simply pattern recognition and does not imply any underlying process. Moreover, comparative data sets generally do not include information that would allow rigorous inferences concerning causal processes underlying such patterns. The concept of phylogenetic inertia needs to be defined and studied with as much care as 'adaptation'.