 There are two obvious answers to this question—two different ways of characterizing the long-term survivors and beneficiaries of the evolution by selection process. One might say that the species or lineages (the previous “evolvers”) are the ultimate beneficiaries of the evolutionary process. Alternatively, one might say that the lineages characterized on the genic level, that is, the surviving alleles, are the relevant long-term beneficiaries. I have not located any authors holding the first view, but, for Richard Dawkins, the latter interpretation is the primary fact about evolution. To arrive at this conclusion, he adds the requirement of agency to the notion of beneficiary (see Hampe &amp; Morgan 1988). A beneficiary, by definition, does not simply passively accrue credit in the long term; it must function as the initiator of a causal pathway (Dawkins 1982a,b). Under this definition, the replicator is causally responsible for all of the various effects that arise further down the biochemical or phenotypic pathway, irrespective of which entities might reap the long-term rewards (Sapienza 2010).