Opening the species box: what parsimonious microscopic models of speciation have to say about macroevolution.

In the last two decades, lineage-based models of diversification, where species are viewed as particles that can divide (speciate) or die (become extinct) at rates depending on some evolving trait, have been very popular tools to study macroevolutionary processes. Here, we argue that this approach cannot be used to break down the inner workings of species diversification and that "opening the species box" is necessary to understand the causes of macroevolution, but that too detailed speciation models also fail to make robust macroevolutionary predictions. We set up a general framework for parsimonious models of speciation that rely on a minimal number of mechanistic principles: (a) reproductive isolation is caused by excessive dissimilarity between genotypes; (b) dissimilarity results from a balance between differentiation processes and homogenizing processes; and (c) dissimilarity can feed back on these processes by decelerating homogenization.