This note provides a definition of evolution by natural selection that is compatible with the extended evolutionary synthesis.

Both biologists and philosophers often make use of simple verbal formulations of necessary and sufficient conditions for evolution by natural selection (ENS). Such summaries go back to Darwin's Origin of Species (especially the "Recapitulation"), but recent ones are more compact.1 Perhaps the most commonly cited formulation is due to Lewontin.2 These summaries tend to have three or four conditions, where the core requirement is a combination of variation, heredity, and fitness differences. The summaries are employed in several ways. First, they (...) are often used in pedagogical contexts, and in showing the coherence of evolutionary theory in response to attacks from outside biology. Second, they are important in discussions of extensions of evolutionary principles to new domains, such as cultural change. The summaries also have intrinsic scientific and philosophical interest as attempts to capture some core principles of evolutionary theory in a highly concise way. Despite their prominence, both the proper formulation and status of these summaries are unclear. Standard formulations are subject to counterexamples, and their relations to formal models of evolutionary change are not straightforward. Here I look closely at these verbal summaries, and at how they relate to formal models. Are the summaries merely rough approximations that have no theoretical role of their own? Perhaps they could operate as theoretical statements in Darwin's time, but have now been superseded by more exact treatments. (shrink)

A lesson in evolutionary theory can be drawn from the work of Rick Charnov, who transformed Fisher’s sex ratio theory into sex allocation theory, but unfortunately, the lesson has not spread far enough. The lesson is that costs as well as frequencies need to be included. That is so whether we are talking about evolutionary ecology (e.g., density dependence), social evolution (e.g., sexual selection), origins, an extended evolutionary synthesis, multilevel selection, or whatever. The two dimensions can be expressed in a (...) variety of ways—investing as well as spending, quality as well as quantity, costs as well as frequencies, and somatic as well as reproductive functions, for example. Both are needed everywhere in evolutionary theory. (shrink)