We look to mitonuclear ecology and the phenomenon of Mother's Curse to argue that the sex of parents and offspring among populations of eukaryotic organisms, as well as the mitochondrial genome, ought to be taken into account in the conceptualization of evolutionary fitness.Subsequently, we show how characterizations of fitness considered by philosophers that do not take sex and the mitochondrial genome into account may suffer. Last, we reflect on the debate regarding the fundamentality of trait versus organism fitness and gesture at the idea that the former lies at the conceptual basis of evolutionary theory.1. Introduction. The concept of "fitness" is central to the foundations of evolutionary biology, especially as it ostensibly plays an explanatory role in how natural selection accounts for much of the diversity and adaptation of the biological realm (Orr, 2009;Rosenberg & Bouchard, 2015). Famously, Herbert Spencer coined the phrase "survival of the fittest" in the 19 th century after reading Charles Darwin's On the Origin of Species and, upon the suggestion of Alfred Russel Wallace, Darwin adopted the phrase into the 5 th edition of his book in 1869.Ronald Fisher (1958) conceptualized and developed much of the mathematics of fitness and