So, I co-architected the Opera Mini infrastructure. It peaked at a similar number of users (250-300M monthly active users). Sure, Twitter is much more DB-intensive, but transcoding web pages is pretty CPU intensive too, and typically we transcoded every single web page for them. Opera Mini was their only browser.
Twitter is spending $5B/300M =~ $17/user per year
I believe that from public sources, it's now possible to deduce that we spent less than a 1/100th of that per user/year, almost a decade ago.
Since we didn't have crazy money, we optimized things at every step. Or, well, mostly avoided doing stupid stuff.