Combining mathematical and empirical work to understand what (if anything) is optimized by evolution across various levels of biological organization.

The view that evolution proceeds progressively to promote the survival of each species is seriously outdated. Still, we lack a hierarchical theory of evolution that quantifies precisely by how much natural or sexual selection fail to maximize any population-level good. Providing such theory combines immense intellectual value with applied consequences for conserving endangered species, limiting the spread of invasive species, combating disease, and understanding the evolutionary roots of our own behaviour. The proposed research additionally builds capacity by importing and spreading cutting-edge knowledge of mathematical modelling techniques that allow integrating genetic processes with their evolutionary and ecological consequences.