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While on four legs, he can be flinch shot into walls.
Not sure if he needs to be standing on two legs to start flying.
The truly important differences are that while Fatalis is on all fours, he is vunerable to wallbangs from flinch shots, as long as he isn't enraged. While bipedal or flying, a wallbang will instead revert Fatalis to all fours and deal much less damage.
Secondly, Fatalis' moveset is different between the stances. His firebreath attacks are a lot more varied while bipedal, but he can charge while on all fours, for example.
Fatalis will transition between these states depending on what attacks he's interested in doing at the moment, but you can forcefully change these states by attacking him. Attacks to the chest while bipedal will encourage a collapse to all fours and attacks to the head while on all fours might make him go bipedal again.
The game's NPCs will talk about hitting Fatalis' head while he's on all fours, and while this is something to note it's not necessarily AS important as they make it out to be. The wallbanging is a much more important consequence of him being on all fours. It's still important to get what head damage in you can before Phase 3, though.
As for how/why to change these states, using the One-Shot Binders (there are two in Schrade, one next to the westmost ballista, and one across to the east side between the cannon and eastmost ballista) will forcefully revert Fatalis back to all fours, and this is especially useful while he's flying. Any Heavy Artillery-subject damage (like the Roaming Ballista, regular ballistae and cannons) also contribute to their own knockdowns versus him which can also be worth landing if his stances are causing trouble.
Aside from this, like most monster AI there is too much randomness involved to totally predict Fatalis' behaviour. He might react to you running away, for example, by either charging straight for you, or sit back upright and unleash loads of fireballs. There's no formula for knowing exactly when Fatalis will start flying.
Thanks for the incredibly detailed answer! The fight feels like it makes a lot more sense now, especially concerning flinch shots.