I made some experiments creating my own bifolios and mimicking the process that I suggested. It appears that this is all a bit complicated.
Supposing that the quire marks are left after the quire binding (stitching) was done, three things lead to the shift of the written part in the upper folio in respect of the part in the lower folio:
1) The binding loosens (or the quire is rebound), leading to the inner bifolio shifting sidewards from the outer bifolio.
2) The binding loosens (or the quire is rebound), leading to the inner bifolio shifting downwards from the outer bifolio.
3) The upper folio gets excessively contorted (e.g. due to the aging)
If all three factors act at once, their effect is accumulating to the same direction.
Clause 2) will additionally leave an y-axis gap between the two parts (we observe the y-axis gap in Q1 and Q3 marks (but not in Q15)).
However, in all three cases the upper part will shift towards the binding. This is what we see in Q3 and Q15. In Q1 the shift is in the opposite direction - outwards from the binding. I was not able to explain this so far. Maybe there's a local downward curvature of f8 in this place.
Supposing that the quire marks are left after the quire binding (stitching) was done, three things lead to the shift of the written part in the upper folio in respect of the part in the lower folio:
1) The binding loosens (or the quire is rebound), leading to the inner bifolio shifting sidewards from the outer bifolio.
2) The binding loosens (or the quire is rebound), leading to the inner bifolio shifting downwards from the outer bifolio.
3) The upper folio gets excessively contorted (e.g. due to the aging)
If all three factors act at once, their effect is accumulating to the same direction.
Clause 2) will additionally leave an y-axis gap between the two parts (we observe the y-axis gap in Q1 and Q3 marks (but not in Q15)).
However, in all three cases the upper part will shift towards the binding. This is what we see in Q3 and Q15. In Q1 the shift is in the opposite direction - outwards from the binding. I was not able to explain this so far. Maybe there's a local downward curvature of f8 in this place.

