Answer: The cultural object or being, whatever it is, not the biological. Dawkins called it a meme and the name’s stuck around, but it is no longer associated with a useful idea. When Dawkins originally proposed the idea in The Selfish Gene he wasn’t sure whether the proposed entity was a thing in the external world, or a thing in the mind. And perhaps it was both. Are we talking about the tune that everyone can hear, a public entity, or a tune that exists in someone’s mind (and which is conveyed to other minds via a pubic entity).
Dennett decided it must be a thing in the mind, he was taken up on it, and the game was lost.
Dual inheritance theory is becoming popular these days. Humans inherit behavioral traits via the genes and via social learning. But, and here’s the point, the evolutionary dynamic is conceptualized as being centered on the biological human, generally a human in a social group. That’s what differentiates it from Dawkinsian memetics. Dawkins got that one right. The others, well...it’s not that culture doesn’t benefit biological human individuals. Of course it does. It has to, otherwise it couldn’t happen much less thrive. But the specifically cultural evolutionary dynamic is operating on the cultural entity, the tune.
As for the tune, there’s a bunch of gene-like entities out there in public space, and there’s a phenotype-like entity in (collective) neural space. But I’m not going over that here and now. It’s all over the place in the blog and in my working papers (e.g. here, https://independent.academia.edu/BillBenzon).