The eco-evolutionary dynamics of such systems have been studied in depth in Takeuchi and Hogeweg (2009), where it was shown that travelling waves constitute a higher level of selection than that of individual replicators and parasites.
The study of the system of differential equations presented in section 2.4 shows that the changes produced by natural selection in the internal population structure of a quasispecies, i.e. entry into error catastrophe, can modify the result of natural selection on the immediately higher level of selection, that of competition between quasispecies.
Even such previous allies as EO Wilson have embraced the possibility of higher levels of selection, of groups and species, leaving Dawkins somewhat isolated in his reductionism.
This distinction between the two meanings of adaptation also turns out to be pivotal in the debates about the efficacy of higher levels of selection, as we will see in sections 3.1 and 3.2.
As Williams himself has acknowledged, in a discussion on species selection: "The answer to all these difficulties must be Lloyd's…idea that higher levels of selection depend, not on emergent characters, but on any and all emergent fitnesses" (1992, p. 27).
Advocates for higher levels of selection (such as Richard Lewontin, David Sloan Wilson, and Elliott Sober) suggest that there are many phenomena (including altruism) that gene-based selection cannot satisfactorily explain.
In fact, we did preliminary runs with a higher per generation change in optimum (e = 0.15), which resulted almost exclusively in extinctions, in particular at higher levels of selection, but also in the other cases.
Consequently, one could also expect that higher recombination rates would inflate ωPAML estimates [60] [62], but that does not seem to be the case, since subtype B showed significantly higher levels of selection than CRF01_AE for all estimators.
Therefore, variability in efficacy of the more abundant components will have the greatest fitness consequences for the snake, and those components should experience higher levels of selection.
If indeed TEs speed evolution, it is at these higher levels of selection that such a function might best be attributed to them as a class.
The importance of this class of genes in plant genomes is also demonstrated by the unusually high level of selection found at disease resistance loci [20], [21].
