A lot of gear discussion, but no matter what kind of tank (or tray), these are development fundamentals of agitation, for me:
Always random in direction - directional patterns in uniform areas ( like your skies) usually indicate a linear direction to agitation. If I'm doing sheet film, open tray, shuffling through top to bottom, rotating the stack each time 90˚. For roll film, in cylindrical tanks (Nikkor, etc), agitation is always inversions, with axis twisted, to avoid patterns. Even then then, I can get edge surge, from the rebate area.
Your artifact, noticing the direction of overdevelopment (the negative image is useful, for me), makes perfect sense to me as bromide drag. Thin areas (shadows, like the trees) do not exhaust the developer. The heavier (sky) areas do, so if the film is moving in the down direction, relative to the trees, there will be relatively unused developer coming into the sky, where the developer is already more used, or exhausted, and will go for the more highly exposed silver halides in the sky. You see them more against the sky areas that are not adjacent to trees, where there is no unused developer entering.
For what it's worth, medium format film has always been most susceptible, to me - more open areas for this to happen in the larger (than 35mm) frame. With sheet film, open tray, the agitation is totally random.