Yep reflexes is a good shout. I also think the ability to adapt during a fight and also to overcome things other fighters can't. Like continuing with injuries, getting up from a painful body shot even if your body doesn't want you to.
I would say it’s more mental than physical advantages in most cases. Pretty much all ATGs have an extremely high ring IQ.
A similar question gets asked in chess all the time. "What do 2200s have that 2000s not?" Or 2500s vs 2300s, 2100s vs 1900s, 1800s vs 1600s, whatever. The answer is that on average, the higher rated players are better at every aspect of the game on average. It's the same for ATGs vs non-ATGs. On average, compared to those of their era, they're better at everything.
I think intelligence always gets overlooked. Not only the in-ring IQ and ability to process data and adjut but also the ability to calm ones mind down and relax in tight situations. The lower level fighters do the opposite. Never really thinking and always tense.
Yeah and it's not just ring IQ, it's the ability to perform when the pressure is on and having the mental fortitude to not crumble under the pressure but rise to that pressure. You see plenty of very talented athletes look unbeatable all year then the championships come around and they just don't perform. Great athletes, including boxers rarely don't show up mentally, no matter what's happening outside the ring once the step inside the ropes they flick a switch and just have laser type focus and unshakeable mental fortitude.
tough question. They fought at high levels where their level is just higher on a consistent basis. Vague answer I know.