 Systems biology is often defined as an approach primarily concerned with functional questions (Boogerd et al. 2007). Evolutionary systems biology (ESB) explicitly seeks to expand this focus through investigations of the evolutionary dynamics of interacting networks (Soyer 2012). ESB aims to put the results of systems biology in a proper evolutionary context and also to enhance and extend current evolutionary biology. Specifically, ESB aims to develop more detailed models of genotype-phenotype maps to study causal changes over evolutionary time-scales and to enhance the predictive potential of evolutionary models. These aims are pursued by combining methods from molecular and evolutionary biology with in silico evolution, comparative genomic analysis, as well as with dynamical modeling of network architectures (Bapteste and Papale 2021; Soyer &amp; O’Malley 2013).