I think it's a server (or collection of servers) that's running redis or something like it. It caches everything posted to whatever relays it's monitoring, and delivers it to the user on request. But posting new notes or replies goes through a user's defined relays.

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Are signatures still validated client side, or can the cache tamper messages?

That I cannot say. I'm deducing its methodology based on what we can see as users. I suspect it may not be preserving signatures, but I just don't know.