Phylogram instead of chronogram when assessing the neutral evolution of a trait

Abstract

Comparative analyses of trait evolution aim to uncover the different evolutionary forces shaping phenotypic diversity among species. This is typically done by fitting various evolutionary models to observed trait changes along a species tree. For instance, under neutral evolution, trait values are modelled as changing randomly along the branches of the tree. In contrast, for a trait under selection, species are typically assumed to track an optimal trait value, which itself may shift along the tree. Here, rather than relying on alternative models to discriminate among evolutionary scenarios, we focus on the underlying species tree, and specifically the units in which its branch lengths are measured. Species trees are usually represented as chronograms, with branch lengths proportional to time. The rationale is that time correlates with trait changes through its connection to the number of generations. However, since the generation time of species can also vary along the phylogenetic tree, we argue that chronograms introduces biases. In contrast, phylograms with branch lengths in units of sequence divergence will account for the effect of changing generation time. In this study, we develop a method to test whether a phylogram provides a better fit than a chronogram for modelling trait evolution. We show using simulations that, for a trait evolving neutrally, the fit of a random evolution model has more support on a phylogram than on a chronogram. However, comparing models and testing different scenarios of selection using a phylogram leads to incorrect predictions. Given these results, we argue that we should use phylograms instead of chronograms when assessing the neutral evolution of a trait. Nevertheless, we support the fact that we should generally continue to use chronograms to model selection acting on a quantitative trait. Lay summaryWhen we look at different species, we can see that some traits are different between them. For example, the brain size of different primate and human lineages varies. The question we ask is: does the time that passed explain the differences in their traits? Or are genetic differences between species better at explaining the differences? The standard in evolutionary biology is to use time to explain differences in traits. However, if the trait is not under natural selection and instead is changing due to accumulating mutations, the genetic differences between species instead of time should best explain the differences in traits. As a result, we argue that using both genetic differences and species divergence time simultaneously provides a better understanding of how traits evolve across species.