   Optimality models put the contingencies of history and developmental and genetic constraints in to the background so as to focus on natural selection. Much work in evolutionary game theory embodies this approach. Most game-theoretic models are used to describe the evolution by natural selection of strategic behavior to a local optimum called an evolutionarily stable strategy (Maynard Smith 1982) (but see Orzack and Hines 2005 and Huttegger 2010 among others). This approach has the assumption that there is direct selection on the trait under analysis. This is often justified on the grounds that there exists a “good fit” between the trait and its current function (Maynard Smith 1978) or that natural selection is ubiquitous in nature (Mayr 1983). The Spandrels paper targeted these general arguments for optimality models.