Wings are too static during the run, I thought it was a hard cloth covering like you see on old medieval paintings! I recommend making the wings undulate a bit or twitch some while the run happens.
The transition between run and fly is also way too sudden, one frame of them ducking down before taking off could help alleviate that.

Too much of a greasy look to it, makes him look like a blobfish. Try to play with colors a bit, reduce and rework contrast some

Widen the jaw some! Males tend to have more squared jaws and more pronounced noses/browlines. Try to play with that a little bit!

I think the shelf is too static now, so I did a thing:

And here's one where I halved the duration on a couple of the recoil frames!

Post the original size, I've a few tweaks to pop on thete!