It is intriguing that not one comment (here or on the site itself) actually addresses the technical content of the article. It's all foaming at the mouth about

It is intriguing that not one comment (here or on the site itself) actually addresses the technical content of the article. It's all foaming at the mouth about Dawkins. The name 'Nowak' rang a bell, so I dug through my citation manager. His papers that I had stocked away are primarily evolutionary game theory in various mathematical settings (static graphs, lattices, or simply aggregate, unstructured populations). He dresses it up in various cute ways, but they're still textbook exercises in game theory with no biological relevance. The argument from Dawkins is really simple: there are heritable traits (call them genes; forget DNA). We need a word besides just "trait" because of transmission of recessive characteristics. Naively, you might say that if an organism has more offspring on average than other organisms in the same population, then there will be more copies of the gene. That's true, but it's not an adequate accounting. The genes of a rabbit that has fifty times as many babies, but they're all runts and die, the genes of that rabbit are not going to show up in the population in the longterm. So you have to talk about the expected long term frequency of the rabbit's genes in the population. And once you realize that, you start looking for all the copies of the gene. It doesn't matter whether the gene in a particular organism is identical by descent or identical by allele. Thus your bookkeeping has to include other paths of descent. However, genes aren't traits (the whole recessive transmission thing). Thus any genetically encoded strategy can't depend on the genes of any other individuals, only on proxies. But proxies can be faked. Among proxies, relatedness is 1) hard to fake and 2) universal among sexually reproducing organisms. Thus it's a really important piece of bookkeeping to do when calculating out expected frequencies of genes. And that's kin selection. Not that it's the only proxy. It's just a really, really common one. As for group vs kin selection, ideas of superorganisms, or such things, it's all a red herring. In the end, what I actually need is a theory describing masses of individual creatures out being born, eating, mating, and dying. Doesn't matter what abstraction you want to couch the case in question in initially, you eventually need to get back to that.