As the Enola lawyers wrote, the generic character traits (their age, emotional vibe, etc) are not protected. Neither is anything "trivial" or "minuscule" - for example, if a cartoonist makes a minor alteration to the way a character's pupils or eyes are drawn, that's a minor detail, not a copyrightable element.
The biggest impediment to using public domain characters isn't copyright, it's trademark.
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