Nathanael is interested in evolutionary novelty, particularly in complex biochemical traits. He investigates the contribution of various sources of variation, especially gene duplication, to evolutionary innovations, using phylogenetics, comparative genomics and gene expression analysis. He is particularly interested in synthesising systems biology and co-expression network techniques to understand how complex traits evolve. He is also interested in core phylogenetic theory, including gene tree conflict, phylogenetic information content, and non-homogeneous evolutionary models.