Ronald Fisher and group selection.

Ronald Aylmer Fisher (1890-1962) was a pioneer of evolutionary biology who founded modern statistics. He has often been associated with a gene-centric conception of natural selection, but he did not discount the importance of factors operating at other levels. In the later chapters of his 1930 book, The Genetical Theory of Natural Selection, Fisher proposed a mechanism to explain how a human civilization of any ethnicity could rise or fall. In contemporary terms, he did this by applying a concept of group selection, also known as "multi-level selection 1", in which supra-individual collectives impart consistent population structure over time to reproductive entities therein. Fisher believed that socioeconomic factors, operating above the individual level, could bias reproductive patterns and thereby have a causal influence on human social complexity. "Multi-level selection 2" is another kind of group selection which asserts heritable features to units above the level of the individual, and Fisher regarded "sexuality itself" as one such example. Fisher held some inaccurate views about human diversity, but appreciating how his argument foreshadows current multi-level selection theory does not require agreement with his mistakes. The chapters concerning human civilization in The Genetical Theory were not a polemic against non-Europeans, but reflected an understanding of multi-level selection and its effects on evolution as a whole.